pteilcl iiin–mliligwtotlsn touch losing without Ste multiplying – division cell Epithelial COMMENTARY ß opsto ftasebaepoen n lipids. by 2 al., deciphered and et are 1 (Apodaca either that machinery to signals trafficking proteins intracellular sorting localized specialized on specifically the distinct transmembrane depending of are a domain, domains have proteins of basolateral domains Transmembrane the These and membrane. composition junction apical plasma adherens the epithelial in an Shotgun between form they and this belt Thus mammals in . in junction actomyosin CDH1 of as type Drosophila known E- this tissue by (also stabilized on maintaining and formed focus in are involved junctions will Adherens are Commentary. we and junctions and Adherens role Tepass integrity, 2001). adhesive 2003; al., basal an Balda, the et are and serve Tepass that invertebrates, (Matter 1994; junctions 1) Hartenstein, in (Fig. and septate junctions by are whereas adherens ensured and to junctions, is barrier barrier diffusion adherens diffusion the to as In cell. act apical the junctions and to tight polarity basal septate intrinsic and vertebrates, an apical or conferring the In between thereby tight limits membrane, contacts. the within cell–cell define junctions various junctions tightly adherens through is intercellular these cell neighbours each particular, multi-layered, its and or to organized, Mono- spatially bound environment. are tissues the between compact barriers and chemical body and the mechanical as function Epithelia Introduction Cytokinesis junctions, Adherens cells, Epithelial WORDS: cell KEY of site the transmitted becoming ultimately is new cells, polarity signalling. where daughter cell remodelling, to and intense mother assembled from of are site contacts the adhesive is interface cells. daughter new during between generated and This is role interface essential cell–cell an new a plays which in belt phenomenon continually junction focus multiparty adherens a will the mitosis, are we which of step Commentary, junctions final this the cytokinesis, In these on division. integrity, cell actomyosin during a epithelial remodelled is the life, junctions as for adherens largely of adult persistence needed to regenerating, Although division. or throughout cell of linked growing result and continuously are proteins development epithelia E- During by stabilized cytoskeleton. and and contacts formed adhesive are on cadherin which part, junctions, in adherens relies, that within barrier cells This and body environment. the juxtaposed between the barriers comprising chemical and mechanical tissues as function compact are Epithelia ABSTRACT Ato o orsodne([email protected]) correspondence for *Author France. Rennes, BIOSIT, SFR UEB, eteNtoa el ehrh cetfqe M 20 ens France. Rennes, 6290, UMR Scientifique, Recherche Universite la de National Centre 04 ulse yTeCmayo ilgssLd|Junlo elSine(04 2,52–17doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal | Ltd Biologists of Company The by Published 2014. pai eBras Le ´phanie ´ eRne ,Isiu eGe de Institut 1, Rennes de n lseso aei rtista r ikdt the to linked are that proteins catenin of clusters and ) 1,2 n oadL Borgne Le Roland and ´ ne ´ iu tDe et tique ´ eopmn eRennes, de veloppement 1,2, * eoelddrn eldvso nodrt lo proper allow to are order junctions in adherens division how on cell light during shed remodelled to begun recently aln 00 ulo n eut 03;Hrzege l,2014). al., et Herszterg 2013b; and Lecuit, Friedl and E- 2011; throughout Guillot and Georgiou, 2010; through formation and Zallen, Baum junction integrity part in to (reviewed in contributes maintenance epithelial which morphogenesis, turnover, continually tissue cadherin barrier are for during junctions mechanical adherens remodelled needed the adulthood, and adherens and development of is persistence polarity the cell Although junctions . global the of the function to Mostov, both and and (Martin-Belmonte 2011). establishment al., polarity et the Shewan cell to 2008; contributes of the along and are support polarized maintenance axis data is lipids phosphoinositides apical-basal elucidated, of fully the which distribution yet the through not that are idea mechanisms cells in the trafficked Although 2012). pteilcl iiin orsuisin during studies contacts Four adhesive division. been of cell have transmission the regarding epithelial cytokinesis known molecular and is little control the cells, maintenance isolated Although that of studies integrity. in mechanisms identified two tissue and the maintain membrane factors to formed between newly order this membrane on in appropriate established new be and must junctions polarity a cell adherens to Furthermore, of order of cells. in formation extent polarity daughter some the to the and remodelled be belt allow to junction need cell adherens cell unknown. dividing the the are epithelial time, changes cells same cell neighbouring the during these At how on and applied impact context, and/or accommodate tissue are a Whether within tensions ingression. cytokinesis furrow cortical allows cortex similar This polar contraction. the undergoes of equator softening the the whereas at relaxed, tension is a – into poles asymmetric exhibit entry become and upon tensions round but cortical are prometaphase, cytokinesis, during cells cortex cells, actomyosin isolated Baum, stiff and of Lancaster cultures 2009; In Baum, by and 2014). (reviewed Kunda 2014; cytoskeleton various al., own to et its Cadart subjected cells. through is exerted sister are cell that dividing forces the the Therefore, two separates the intercellular generating cells. thereby at distinct midbody, narrow bridge, the membrane the a resolves eventually plasma Abscission by until the stabilized furrowing) of the bridge, cleavage then ingression generate this (i.e. local then constriction; equator the ring II) for to (Myo needed 1). leads is (Box II that equator myosin tension cell contractile is the non-muscle that at and ring membrane contractile plasma actomyosin the to an connected of assembly the mediates 2012; and al., (Fededa et division 2004). Green Spudich, cell 2010; and Glotzer, of Robinson 2005; stages Glotzer, later 2012; is the Gerlich, which during integrity cells), during two particular tissue initiated into in separates how is cell divides, mitotic to the cell regard (when epithelial cytokinesis with an known when is maintained little so, Even naleihla el,teahrn ucinbl contributes belt junction adherens the cells, epithelial all In olwn itrcrmtdsprto,teaahs spindle anaphase the separation, chromatid sister Following Drosophila 5127 have

Journal of Cell Science COMMENTARY hycndvd ihu eknn h are ucin Several 5128 function. barrier the weakening how without of question divide the can raises this they to divide, connected they mechanically while remain neighbours epithelium their an in cytokinesis? cells epithelial platform Because during polarity signalling of Maintenance a as behaving cells. the daughter interface to the between leads of membrane and interface, cytokinesis adhesive anchoring resolves new new abscission the the how of finally, furrow, formation during and, the cleavage junctions and adherens the ring of contractile of role cell progression the daughter discuss the the to will transmission we dynamics its the interface; and on maintenance focus polarity Morais-de-Sa will of Commentary 2014; this al., Therefore, 2013b). et (Herszterg in cytokinesis Kotak involved Go 2014; al., forces et and the (Almonacid mitosis discussed during positioning have spindle reviews recent Morais-de-Sa excellent 2013; al., et complexes Herszterg adhesive and markers vivo polarity cell of p as partitioning IV; known Neurexin also Lin7, Nrx-IV: with larvae; associated giant protein (2) PALS: ARHGEF18; Lethal or Lgl: p114RhoGEF moesin; as and known also radixin p114: ezrin, 1994); ERM: Karsenti, sept in E-cadherin; and belt: Stardust (Reinsch E-cad: junction in antigen large; adherens junctions 58-kDa Discs the as adherens Dlg: known to the Crumbs; relative above Crb: and barrier prometap Coracle; below diffusion interphase, Cora: junctions, the during adherens of to insets) position relative the the midbody in is in (listed epithelia apical phosphoinositides these are and between (orange) markers difference polarity fundamental complexes, mitosis. A junctional during cytokinesis. of markers composition polarity and apical-basal distribution of Distribution 1. Fig. ,,-rshsht;Srb cibe O oaoccludens. Zona ZO: Scribble; Scrib: 3,4,5-trisphosphate; Fuonue l,21;GiltadLci,2013a; Lecuit, and Guillot 2013; al., et (Founounou nz,21;MNly 03 n h ehnc of mechanics the and 2013) McNally, 2013; ¨nczy, Drosophila AJ rti soitdwt ih ucin;PtdIns(4,5) junctions; tight with associated protein PATJ: ; Drosophila hra ih ucin prl)aebsli etbae.Amjrdfeec ewe pce stefnlpsto fthe of position final the is species between difference major A vertebrates. in basal are (purple) junctions tight whereas , ´ n ukl 03) Other 2013a). Sunkel, and ´ n Sunkel, and pteilclsin cells Epithelial in P 2 hshtdlnstl45bshsht;PtdIns(3,4,5) 4,5-bisphosphate; phosphatidylinositol : netgtosuigtm-as irsoy npriua,the particular, invertebrates in junctions In septate of instance, components microscopy. the for of localization time-lapse – awaits using still investigations demonstration formal dividing of cells, polarity membrane epithelial apical-basal the in change is global there no that imply collectively studies aforementioned the in although depolarized (Morais-de-Sa partially cells epithelial is follicular it whereas 2013a), a6 uigmtssapast eedo h ellrcnet sit as context, in cellular unaffected the on is depend and to aPKC appears mitosis (Par3/Bazooka, pupal during determinants Par6) in apical Notch the and of Crumbs localization E-cadherin, as (Devenport well Drosophila skin as mammalian 2011), and MDCK al., 1993) in et Garrod, and Reinsch and (Baker 1993; ZO-1 cells , Garrod, apical-basal 1994), and Karsenti, (Baker appropriate and cells Madin-Darby (MDCK) or their in kidney E-cadherin proteins canine e.g. at mitosis, transmembrane throughout remain localization several proteins during Indeed, cells adhesive associated 1). epithelial remain (Fig. that and polarity division suggest apical-basal systems their model maintain different in studies Drosophila Drosophila ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal n etbae,rsetvl.aK:ayia rti iaeC; kinase protein atypical aPKC: respectively. vertebrates, and pteim(onuo ta. 03.B otat the contrast, By 2013). al., et (Founounou epithelium Drosophila lf)advrerts(ih)miti hi apical-basal their maintain (right) vertebrates and (left) mroetdr GiltadLecuit, and (Guillot ectoderm embryo ´ n ukl 03) Thus, 2013a). Sunkel, and P 3 phosphatidylinositol : Drosophila aeand hase t junctions ate 8 also 58: ovary

Journal of Cell Science on nvrertsadi net r ifrn PE versus (PTEN different are PtdIns(4,5) insects apical of in kinases function the and lipid Skittles), the vertebrates Although in 2014). al., found epithelial et i.e. (Claret (EMT) constriction, transition apical epithelial–mesenchymal the of in reminiscent changes results morphological that actomyosin active ring of apically disassembly contractile an the of formation causing and thus junctions Par3, adherens determinant polarity the of el,a ihihe npeiu eiw Bile l,2011; al., et (Brill reviews isolated previous of in cytokinesis highlighted throughout as remodelling trafficking cells, cytoskeletal membrane in roles and major have phosphoinositides conserved. evolutionarily hc ntrsrcut nei n h Cdc42–Par6–aPKC Reciprocally, the 2007). and al., 2 et phosphatidylinositol (Martin-Belmonte annexin (3,4,5)-trisphosphate complex [PtdIns(3,4,5) recruits polarity turns in which PE) nue h pclsgeaino PtdIns(4,5) of segregation apical homolog tensin the and induces apically phosphatase the phosphatase, (PTEN), polarization, cell lipid epithelial localized During 1). (Fig. surface aoaea PtdIns(3,4,5) basolateral COMMENTARY el rw scssrvae htpopaiyioio 4,5- phosphatidylinositol that revealed cysts [PtdIns(4,5) as MDCK mitosis bisphosphate of during grown Observations proteins phosphoinositides. cells of polarity that of the cortical mirror localization would on the the might relies that one As above phosphoinositides, predict proteins. discussed specific less of markers polarity even membrane-localization polarity of is of that division localization than cell epithelial characterized during the phosphoinositides from thus barrier unexplored. and largely diffusion remains mitosis, cells, the daughter during of to mother vertebrates maintenance and/or in transmission junctions tight or ktlsrdcsPtdIns(4,5) reduces Skittles the of DKcss(asm-igee l,20) In PtdIns(4,5) 2006). in also, al., membrane basolateral epithelium et the (Gassama-Diagne follicular of cysts formation the MDCK regulates and to ufc n PtdIns(3,4,5) and surface h rdei vnulyrsle,teeygnrtn w distinct two generating thereby the resolved, form cells. eventually to is matures bridge abscission, midzone Upon the bridge. spindle intercellular the the 2). organizes (Fig. which constricts, and midbody, constriction to ring ring bind filament to the that leading the As filaments, filaments actin bipolar and on forms forces filaments II exert Myo heteromeric anillin. linear linker cross form membrane-associated II, that Myo septins composed filaments, network actin the filamentous formin-nucleated a induces – RhoA of in ring active contractile membrane There, the II. plasma as of equator, assembly Myo the the of at onto phosphorylation RhoA as of it activation well localized loads the and promote factor sister to – exchange order between nucleotide GTPases midzone guanine Rho the a for – in Ect2 forms active recruits of spindle chromatids, zone chromatid equatorial central an sister The generate to RhoA. following cortex cell Briefly, the that anaphase and the spindle mind growing. between signalling in with rapidly starts bearing cytokinesis separation, 2004), is Spudich, Green field and 2010; this Robinson Glotzer, core 2012; 2005; Glotzer, the al., topic this 2012; et on of Gerlich, reviews excellent and identification and previous (Fededa to cells, the refer isolated should in readers to the cytokinesis led underlying machinery have molecular studies Numerous cytokinesis of overview Brief 1. Box nadto oterrl naia-aa polarity, apical-basal in role their to addition In h pclbslpam ebaecmoiinof composition membrane plasma apical-basal The Drosophila hshtdlnstl4popae5-kinase phosphatidylinositol-4-phosphate P P P 3 3 2 ntsu nert per obe to appears integrity tissue in , P sacuildtriato h apical the of determinant crucial a is ] h aoaea ufc.Ide,loss Indeed, surface. basolateral the 2 eesadipisaia targeting apical impairs and levels P P 2 2 n rbbyta of that probably and , eie h apical the defines P Drosophila 3 localizes ] P 2 , adt,20;Nzse l,21) oee,terrl in role PtdIns(4,5) their of for However, segregation unclear, is apical-basal 2010). it whether and and al., example, unknown, Logan remains et cytokinesis 2006; cell Nezis epithelial Devreotes, 2006; and Mandato, Janetopoulos 2012; Echard, oal eeae ntePtdIns(3,4,5) the in generated locally PtdIns(3,4,5) motn oe nctknss hsrie h usino how of question the raises exert plasma This interphase, apical cytokinesis. PtdIns(4,5) during in the cells roles epithelial at important polarized enriched in are membrane which glycosphingolipids, nteedt,oecudatcpt htPtdIns(4,5) that that anticipate cortical revealed could controlling Based one cells 2014). in data, al., isolated et role these (Atilla-Gokcumen in on at key mitosis localization during cytokinesis a and stiffness have composition of lipid glycosphingolipids onset in approach changes lipidomic the recent the A study 2010). ring al., to Maddox, et contractile D’Avino and trigger 2012; Piekny to al., 2009; et 2) (Chesneau Fig. constriction 1; and assembly (Box Rho for septins (GEF) and factor II, exchange GTPases] Myo nucleotide guanine F-actin, [a including Ect2 cytoskeleton, in RhoA, actin ANLN the as of known in components (also Scraps anillin and of mammals recruitment the for essential urwn’ rcesfo h aa oteaia membrane, well as 2001), apical al., and (Reinsch et ‘asymmetric monolayer the (Kojima a hepatocytes as cultured (or to grown 1994), Karsenti, are that basal ingression cells MDCK in the cell unequal as such epithelial from planar cases, proceeds of type most furrowing’) belt. this junction on In focus adherens will division. the epithelium, we bisects the on, of ring here plane contractile From the the to when parallel apical-basal divides i.e. the initially cell along Similarly, the an furrow 2012). when cleavage by axis al., the characterized of et ingression generally Tse unequal is the 2007; cytokinesis al., of epithelial et that (Maddox is ‘asymmetricfurrowing’ promoting mechanism, example symmetry-breaking the a physiological follows of For more ring and A contractile rule. (Fishkind substratum asymmetric 1993). the the is to Wang, furrowing be adhered division, remain cell not that throughout cells manner, might isolated symmetric in furrowing 2013). example, a al., symmetric in et (Founounou contract actomyosin However, ingression’ typically the furrow bisect ‘symmetric to cytokinesis, plane not termed assumed during the does is ring belt to contractile ring furrowing junction orthogonally the adherens when divide is isolated i.e. the that how epithelium, In cells the controlled? of of epithelial spatially question or cells cells epithelial the furrowing the in raising in progression probably, thus very differs and, composition membrane, axis membrane apical-basal plasma the progression the along furrow above, cleavage described in junctions As adherens for role A ebaeo h laaefro.Tee PtdIns(4,5) plasma There, the furrow. at cleavage enriched the obviously of is membrane that phosphoinositide nciaegn rdcsi eie eprladsailmanner 2012). spatial al., and in to temporal et able defined (Idevall-Hagren mitosis a are in that during products tools gene glycolipids optogenetic inactivate with or example, lipids for combination, dynamic specific the of of detection allow fascinating localization that these probes reliable Addressing requires issues initiates? furrowing where domain ta. 2000), in al., et that, established well 05 ornie l,20) PtdIns(4,5) 2006), al., al., et Field et 2005; al., al., et Kouranti et Emoto (Abe 2011; cells 2005; al., mammalian et isolated Dambournet and 2012; 2011) al., et Roubinet ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal P 2 P 3 Drosophila n lcshnoiisaetree oand/or to targeted are glycosphingolipids and speevddrn pteilcl ioi.I is It mitosis. cell epithelial during preserved is anradtselegans Caenorhabditis Drosophila 2cls(e lKdie l,2011; al., et Kadhi El (Ben cells S2 cioacaoye pombe Schizosaccharomyces ,wihte neat with interacts then which ), P 3 erce basolateral -enriched P 2 yoe which zygote, steonly the is P (Zhang P 2 P 2 5129 2 and and is

Journal of Cell Science COMMENTARY opsto ftecnrciern,wt eadt y II, Myo to 5130 regard basal (Founounou with cytokinesis versus throughout ring, apical changes Anillin, contractile the and the septins that of revealed ring has composition In contractile analysis 2007). asymmetric al., time-lapse an et 2011; (Maddox organize al., structure which et Anillin-1, Page and Le 2013; and the In Guillot al., 2013; et al., that et Morais-de-Sa cases Herszterg (Founounou the to 2013a; far of shown so most Lecuit, been in reported has ingression been furrow have positioning unequal apical from 2013), furrowing result additional asymmetric al., apical of achieved et Although absence be the the also (Herszterg 2). in can extrusion apically in 1, apical midbody (Figs through the results midbody positioning of basal-to- the ingression means This of 2013). furrow al., positioning et notum asymmetric pupal Herszterg and 2013; apical 2013a) al., recently, Lecuit, et and (Founounou More in (Guillot 2004). observed ectoderm (Morais-de-Sa embryo al., been epithelium et also follicular has ovary Kosodo furrowing 2007; al., asymmetric 2003; al., et et (Das Dubreuil neuroepithelium vertebrate and 1992) Ishikawa, as link which (green), E-cadherin as il such Schematic molecules, (i) adhesion 2013). by a al., mediated is et and constriction (Founounou junctions initial Elsevier adherens the from by middle, permission through supported the with are cytoskeleton in reproduced that actin notum Founounou; cells pupal N. between the of contacts of courtesy adhesive images Image the constriction. in anchoring. ring shown ring ( contractile As and ring phase. progression contractile furrowing furrow the Asymmetric and cleavage (A) cells. in epithelial junctions in adherens progression of Role 2. 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The furrowing. of symmetric courtesy and Image Anillin, and constriction. septins ring II, contractile Myo of of distributions start polarized the the of disappearance the with coincides MyoII ctnnado h idn fMoI iiflmnst -ci.A h nta pcladbslrtso otatl igcntito ifradcor and differ constriction ring contractile of rates basal and apical initial the As F-actin. to mini-filaments II Myo of binding the and/or -catenin nvivo in :yI–Cer,rd rmtebl fahrn ucin Ecdei–F,gen rohaspitt h paetrn eahet.Ti step This detachment). ring apparent the to point arrowheads green; (E-cadherin–GFP, junctions adherens of belt the from red) ::MyoII–mCherry, .elegans C. nmueitsie(lmn ta. 07 igj and Jinguji 2007; al., et (Fleming intestine mouse in ´ n ukl 03;RishadKret,1994). 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Drosophila a ctnni motn o eriigadognzn ehoko yokltlpoen,icuigF-actin including proteins, cytoskeletal of meshwork a organizing and recruiting for important is -catenin ´ n ukl 2013a), Sunkel, and pteilcells, epithelial Drosophila oaie deesjntos(ulo n eut 2013a; Lecuit, apically and with (Guillot associated junctions closely Morais-de-Sa adherens remains the Interestingly, localized cells? ring epithelial these Herszterg contractile in in 2013a; and achieved Morais-de-Sa Lecuit, asymmetry ingression and 2013; are septins Guillot al., they furrow that 2013; et contraction, al., fact unequal polarized et actomyosin the (Founounou of for transient rate and Morais-de-Sa dispensable the the factors 2013a; regulate despite these Anillin Lecuit, of However, and distribution 2013a). 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Journal of Cell Science raie yahrn ucin n iswti hi plane their within is lies one and first junctions The adherens coexist. distinct by two belt, meshworks organized adhesive the actomyosin of belt level contractile junction the adherens at the the cytokinesis, to During ring how contractile the adherens Anchoring of the bisects question ultimately and the belt. to junction anchors raising ring contractile furrowing, asymmetric COMMENTARY agtritrae oee,tedvdn eli ne the under in studies is multiple recent to cell Although subject thus forces. dividing and tensile neighbours its the of constraints daughter– However, physical nascent the the interface. at before complexes daughter furrow E-cadherin cleavage between new the of complexes at assembly cells E-cadherin neighbouring of and cells mitotic disengagement daughter between the junctions adherens requires new of formation interface The adhesive new new a Generating of formation synchronized, below. and address the will orchestrated we and is which complexes furrowing how junction of between question adherens the coupling raising generated, the is cells daughter between al., of et furrow (Founounou 2013). rate belt eventually al., symmetric et junction is Herszterg the that adherens 2013; apparently midbody the a and below in an results positioned composition This 2B). in (Fig. protein ingression resulting becomes the ring constriction, contractile E- in the of 2013), loss homogenous al., localized et ‘apparent’ (Founounou the cadherin of with domains concomitantly initiates basal furrow and asymmetric in apical at participating anchoring separating the thereby ingression. to of ring, of contribute contractile mode properties would the the junctions intrinsic through in adherens the of ring change Anillin level local contractile ring the for This contractile the 2Aiii). the substitute anchors adherens (Fig. the anchor that to belt propose locally we analogy, able junction to to By 2010). al., is order bind et (Goldbach 2008; in 2006), to Nelson, E- septins and Nelson, ability an and/or Hartsock 2013; and septins, its al., and/or et Weis to (Desai Anillin filaments owing of actin the complex, absence in the cadherin–catenin Interestingly, in junctions. anchored cytokinesis) adherens are that by hemi-rings developing separated two constriction. and of of impression rates to II, their the Myo in gives of and above), This distribution protein (see polarized in Anillin is the differ and ring septins by ingression contractile judged the the as furrow ultimately of phase, composition, this domains and During basal 2A), surface midbody. and (Fig. the apical apical of the positioning phase apical to to basal leads first the from the asymmetric epithelial In in that, to phases. revealed able of have the they constriction cells of ring are quantitation and of how analysis parameters Time-lapse composition, 2Aiii)? their contractile (Fig. connect two cytokinetic in these the the differ Because of 2Aii). meshworks plane is (Fig. the belt to meshwork junction orthogonally adherens positioned second is that the ring contractile whereas 2Ai), (Fig. ukl 03) nipratusle usincnen the concerns Morais-de-Sa question 2013; unsolved important al., an et Guillot 2013a), Herszterg 2013; Sunkel, 2013a; its al., and Lecuit, et cell (Founounou and dividing cytokinesis a between during coupling neighbours mechanical the on light ial,a urwn rces e deieinterface adhesive new a proceeds, furrowing as Finally, that progression furrow of phase second the during contrast, By Drosophila Drosophila ua ou,froigpoed ntwo in proceeds furrowing notum, pupal aegrln a xml fincomplete of example (an germline male Drosophila aeshed have a -catenin ´ and edn oadces nEcdei ees oee,actin however, levels; cells, E-cadherin daughter in the decrease between cells in a epidermal interface to neighbouring the the leading of Before symmetric from 2013). during membranes withdraw membrane al., the cell et daughter furrowing, Herszterg 3A) the (Fig. 2013; of belt nascent juxtaposition al., junction the of adherens et sides the both (Founounou with on Myo alignment cells of in adjacent accumulation interface immediately the by the shown in as II cells tension, al., extrinsic neighbouring an constriction, the et exert with ring Herszterg concomitant of Indeed, process 2013; 1992). cell-autonomous Ishikawa, al., and et Jinguji maintained 2013; membranes (Founounou be to of mitosis appears cohesion cells throughout neighbouring epithelium, and mitotic notum between pupal in However, ta. 09.Terneo h uln ocsrqie olocally to required forces to pulling required the force of range the (Zhang The pN contrast, 50– 2009). 50 By around of al., is range et 2004). dimers the or al., monomers in forces E-cadherin et are separate pulling plasma cells (Chu the two nN separate that 200 neighbouring to shown required has are E-cadherin which and/or using express study juxtaposed that A adherens forces. cells membrane the pulling closely for requires to which relative needed membranes, plasma oriented are indeed be to are membranes nascent have forces they two disengagement, tensile later junction a If the at 3B,C). furrow or (Fig. unknown when decreasing, during remains are occurs levels stage, it junctions E-cadherin epithelium, when the notum ingression, of pupal In is disengagement 2013). this al., whether et of activity membrane (Herszterg junction case cell new Myo-II–ROCK-dependent the dividing the of the length the of Indeed, thus mechanical juxtaposition and the a it. for exerting responsible by on interface adhesive force of length formed the control newly actively membrane cells the neighbouring cell the dividing and the 3C), of (Fig. juxtaposition the after minutes few ring. contractile which the of formin in all 2011), localized the al., are RhoGEF2, et in (Levayer of and, ROCK activation and 1998) Diaphanous through al., protein initiated et be (Bauer to cells found mitotic in shown its been occur has to E-cadherin to of due endocytosis support hypothesis, In latter be this membrane. could of ingressing 2002). staining the al., within E-cadherin endocytosis et subsequent in localized (Fujita for decrease decrease the the the through, Finally, for and account E-cadherin could ubiquitylation degradation of Hakai-mediated degradation staining. example, E-cadherin of local of dilution delivery a Alternatively, the in result of would absence E-cadherin, the new in which, membrane is a ingressing within trafficking the First, levels vesicular supply through the posed. components E-cadherin to membrane linked new be be of in could E-cadherin can of decrease amount the hypotheses in reduction the Several of unknown. currently basis 2013). molecular al., et the Herszterg 2013; and The al., interface et (Founounou this 3B) of (Fig. vicinity midbody the in place takes polymerization tteeg ftefro Fg 3B (Fig. neighbours furrow its and the the cell of and mitotic edge the the separation between at ‘hole’ physical a in of formation results are disengagement Lecuit, junction that and the Guillot In 2013; certain forces al., a 2013a). et (Founounou reach cells tensile mitotic neighbours to the between their strength need and adhesive the which the overcome ring, underlie to by contractile threshold the induced by that mediated is mechanisms on disengagement based model, A molecular and junctions. adherens of and disengagement timing olwn hs h omto fnwahrn ucin eisa begins junctions adherens new of formation the this, Following septin ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal uatpeoye,sget htjunction that suggests phenotypes, mutant Drosophila mroi coem hsadherens this ectoderm, embryonic 0 GiltadLci,2013a). 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Journal of Cell Science COMMENTARY elc,21;ArmyradMri-ern,21;Green 2013; 5132 Martin-Serrano, and and (Fededa machinery Agromayor abscission the 2012; role for central Gerlich, scaffold a a have a to forming for proposed in 2 cells, ring been isolated Box have in (see contractile microtubules work ring midbody remarkable midbody and on Based and description). spindle midbody detailed the central form to ring the mature contractile the actomyosin completion, the of second of reaches separation constriction the physical As the and cells. daughter to ring, leading first contractile the abscission, the with comprising steps, of main constriction two into comprising cut subdivided final be the can – Cytokinesis abscission to route En forces will measurement these enabling issue of sensors, this tension Addressing of established. development be the require the to by yet exerted is forces ring pulling contractile the with and compatible unknown, are is they whether cytokinesis during junctions adherens disengage enlarge shown are C and B A, in highlighted zones The junctions. adherens the of plane the within interface. top adhesive the new from a viewed of A are generation insets Cells the neighbours. and its of disengagement context junction Adherens 3. Fig. h egbuigclsta r nue yMoI ntedvdn el(lc rosi C in arrows (black cell dividing the in II Myo by induced are that cells neighbouring the B B in arrow (blue side one In on contacts. cell adhesive dividing the a through in (black connected ring (B forces remain contractile tensile furrow cell of cleavage generation neighbouring the the and at drives C dividing accumulation neighbours green, II its (dark Myo and interface (B). cell furrow daughter–daughter the mitotic of the edge between the complexes at II) A (Myo in II arrows Myosin B (blue of furrowing accumulation of subsequent initiation the the and during cells neighbouring the on deiecnat r vnulyfre dr green). a (dark cell formed mitotic eventually the are between contacts gap adhesive a of appearance the and disengagement junction adherens in in contrast, resulting By interactions, neighbour. homophilic E-cadherin of strength 9 9 nteigesn ebae.Fraino e deesjntosbtenteduhe el eurstedssebyo -ahrn(-a,lgtg light (E-cad, E-cadherin of disassembly the requires cells daughter the between junctions adherens new of Formation membranes. ingressing the on ) ;hr,teeattmn fahrn ucindsnaeeti rsnl nnw.()A urwigeso rcesadtemdoyfrs esl f tensile forms, midbody the and proceeds ingression furrow As (C) unknown. presently is disengagement junction adherens of timing exact the here, ); 9 ,B 9 –B 0 n C and nsitu in 9 epciey A tteaahs-otlpaetasto,teasmldcnrciern rdcrl nA n )eet uln forces pulling exerts C) and B A, in circle (red ring contractile assembled the transition, anaphase-to-telophase the At (A) respectively. , . Drosophila ua ou n os netn,temmrn ftedvdn elrmiscnetdt hto t egbu sonin (shown neighbour its of that to connected remains cell dividing the of membrane the intestine, mouse and notum pupal 9 .I sdpnetuo h ellrcneta owehri h ra urudn h urwmmrnso the of membranes furrow the surrounding areas the in whether to as context cellular the upon dependent is It ). 0 n hs ytenihorcl nteohrsd bakarwi B in arrow (black side other the on cell neighbour the by those and ) 9 ,B 9 n B and Drosophila 9 ean imycoe nebysta akmidbody lack that embryos 2013). al., cells in et daughter closed (Green between mammals) firmly interface in cell–cell remains SPD-1 PRC1 the protein the as bundling midbody Furthermore, that microtubule the known embryos raising the in not of (also seen steps, but depleted as midbody been abscission, itself, both have for ring that crucial midbody are for found microtubules the dispensable also that possibility study are (ESCRT) this transport microtubules for Strikingly, required during and complex machinery. cells septins and sorting on daughter relies midbody endosomal steps the second the the of This division. one abscission), cell into following of the released In process are function. ring septin (the of midbody step absence cells the second in daughter delayed of the is formation cytoplasm this the the isolated; First, be allows 2013). to bridge al., intercellular et been an (Green of recently phases in has two study in view proceeds a accepted commonly in This revisited 2012). al., et n B and 0 9 .Ti ed otelclatvto fRokns ntenihorn cell, neighbouring the in kinase Rho of activation local the to leads This ). lo h ls utpsto fduhe elmmrns nwihnew which on membranes, cell daughter of juxtaposition close the allow ) 0 ,floe yteasml fnwEcdei opee tthe at complexes E-cadherin new of assembly the by followed ), ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal mroi coem h poigtniefre xre ythe by exerted forces tensile opposing the ectoderm, embryonic lutae eei yoiei fa pteilcl ihnthe within cell epithelial an of cytokinesis is here Illustrated .elegans C. mro hr abscission where embryos 0 ih yasteadhesive the bypass might ) rw in rrows dits nd re in orces reen) din

Journal of Cell Science COMMENTARY emieadsmtctsuso aiu pce,including species, various of of the devoid tissues in are ago somatic decades cells and discovered daughter were germline canals epithelial Ring connect microtubules. ring as that to referred 2007). canals, also al., bridges, et that intercellular (Dubreuil belt cytoplasmic midbodies junction Second, maturing adherens the in above detected found that are longer indicate no cells and microscopy are neuroepithelial mouse microtubules electron spatially in First, cytokinesis also of abscission. studies is mouse to in microtubules uncoupled because temporally midbody in nematodes, of to and disappearance unique. specific cells not mechanism neuroepithelial microtubules is redundant midbody cytokinetic for abscission worms, requirement this cell– in a of in stable make lack a the least and of Nonetheless, 2010; formation at interface al., the Thus, to cell et contribute 2005). Estey might al., mechanisms 2011; et Martin-Serrano, Mollinari and mammalian isolated (Caballe in cause cells conditions multinucleation these and whereas 2007), abscission al., defective et (Green Maddox septins or 2013; proteins al., ESCRT et of depleted are or microtubules ta. 01 e ta. 08 oiae l,20;Za ta. 2006). al., et Zhao 2007; al., et Morita membrane 2008; al., Guizetti drive et 2005; and Lee al., 2011; to Carlton et al., Fabbro 2008; 2011; et polymerize al., al., et et and Elia Carlton 2007; 2010; al., Martin-Serrano, 2008) Barr, and et al., the (Bastos (Lafaurie-Janvore scission recruit et they spastin Yang of abscission, 2013; enzyme stages in ESCRT severing functions proteins. later main CHMP4A) ESCRT-III microtubule the two as recruits known have turn in (also proteins in alix complex which to ESCRT-I, spindlin binds and which also central CEP55, CEP55 through the cytokinesis. recruited to is machinery binds human ESCRT isolated In the and 2013). the ESCRT cells, (Agromayor al., et event the McCullough scission narrow and 2013; final Martin-Serrano, 2013) the that promotes Prekeris, which that and components machinery machinery, (Schiel abscission bridge trafficking respective the abscission intercellular of membrane the to platform recruitment a ring includes as the Although serve midbody allows to proposed 2012). and that is midbody midbody the al., unknown, the remains RhoGAP the et of of domain contributions C1 (Lekomtsev the for a mammals, proposed CYK4 In been midbody 2012). has al., the role et of (Kechad similar which membrane attachment plasma the domain, the enables to C-terminal 2012; ring filaments, required its is septin al., II, whereas to et Myo integrity, binds and Kechad ring Actin both midbody 2012; to for binds In al., which et Anillin, 1998). of Hu domain al., 2011; et the al., Madaule of et and filaments components septin (Gai some II, Myo anillin retains filaments, the actin ring as of (Lekomtsev such for midbody ring, membrane contractile The ability required plasma BIRC5) 2012). the is the al., to as et midbody phosphoinositides, through the to known of complex, attachment bind midbody, spindlin (also the to with central MgcRacGAP associated survivin (also stays the the borealin CPC and whereas 2012). and (INCENP), al., CDCA8) aurora et 2012), protein (Carmena as comprising Glotzer, centromere inner and (CPC), known B, White complex kinase mammals; passenger in GTPase RACGAP1 chromosomal family as CYK4 Rho known the (also or and MgcRacGAP (GAP) mammals) Mklp1 protein in the kinesin activating KIF23 the contains as comprising known and complex (also spindlin 2012) Shaw, central and Walczak al., the 2009; 2010), (Glotzer, et PRC1 linker Green cross microtubule 2009; microtubule (Glotzer, antiparallel packed network densely a comprises midbody The composition midbody abscission of and overview Brief 2. Box Drosophila Drosophila 2cls h N-terminal the cells, S2 pteilcls the cells, epithelial aa oiinn,i ohcss igcnl n maturing that be and might forces topology tensile canals This belt. the junction ring from adherens the removed by cases, be junctions induced are to both found adherens epithelia, apical- are final in midbodies the mouse above in positioning, difference junctions this in located basal Despite adherens 2007). whereas al., are et to (Dubreuil 2013), midbodies basal Cooley, found maturing simply and are are canals they (McLean whether ring or are epithelia, function, In abscission. canals cell specific epithelial in a ring intermediates in fulfill cytokinesis whether arrested to temporally to with order These as cells 2013). in has question found Cooley, function exclusively the no and and (McLean raise cycles, them observations cell with two ring associated than of been less capability diffusion for the ensues epithelia canals larval follicular in between cells, equilibration functional protein epithelial allow remain and which days canals, protein Cooley, several ring for and ovarian or McLean to 2011; contrast proteins al., In 2013). et cytoplasmic (Airoldi diffuse specific can complexes some channels intercellular which direct as In act through oocyte. can the canals ring towards tissues, cytoplasm somatic nurse-cell of al., amount et extensive Greenbaum Fawcett, Ventela 1978; 1998; and Drosophila Giorgi, Spradling, and 1958; Dym Pepling Ito, 1980; 2009; and Arnold, Fawcett and 1971; (Cartwright mammals uorgnct rvee yCe ta. 03 cikand Schink 2013; al., et Nonetheless, and Chen 2011). reprogramming Stenmark, by cellular identity, (reviewed cell cell tumourigenicity including regulate stem contexts, to of developmental proposed maintenance various been in have determination that and fate debris cellular are cells epithelial in localization dynamics midbody ESCRT-III the monitor investigate of to to and interesting abscission to be up would leading maturation it 2009), al., trigger et (Vaccari ESCRT-III to in of multinucleation loss required cause As also isolated 2013). is can al., in tension et abscission (Lafaurie-Janvore has and cells in complex study human ESCRT-III reduction recent the a local of recruitment as a abscission that of completion shown the for essential ean otoesa,oe h atfwyasi a eoeclear become has it years determination few past fate the cell over in controversial, remnants remains midbody of interface role membrane the new Although the at signalling Cell–cell ucini ie ellrcnet ssilhgl eae and debated highly still is contexts demonstration. the direct cellular signalling awaits their although given random, conclusion, in not In is function remnants 2014). midbody al., of et partitioning dispensable (Ou be to development appears for cues, that early polarity remnants antero-posterior in in midbody on of cells relies Finally, partitioning adjacent 2014). stereotypical by the al., or embryos, 2012) et interfaces al., for (Crowell phagocytes et in vertebrates by (Chai up occur worms be taken in be to to degradation to medium, found and this cells also extracellular of non-daughter between are spite the In remnants dorso- into 2014). midbody the released Pohl, evidence, of and direct part (Singh latter in mediates formation axis remnants work ventral midbody recent of evidence, positioning correlative are above on elegans outlined functions primarily possible the based Although (Salzmann identity 2014). cell al., of stem dictating et inheritance not asymmetric are for they in rings, example midbody stem least stereotypical a germline at are female Thus, which whereas it. ring, inherit midbody cells the exclude cells h rcs fasiso eut nmdoyrmat that remnants midbody in results abscission of process The ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal mro a ietyrvae htteasymmetric the that revealed directly has embryos eaegrln,rn aasesr h rnfro an of transfer the ensure canals ring germline, female Drosophila Drosophila Drosophila ta. 03.I the In 2003). al., et ¨ Drosophila nvivo in aegrln stem germline male olclrepithelium follicular emielineages, germline . .elegans C. igdisc wing 5133 C.

Journal of Cell Science omncto.Poal n ftebs xmlsi the is examples best cell–cell the biases of and/or one enables Probably a that opens communication. interface, cell–cell window new a spatiotemporal generating by cytokinesis, that COMMENTARY 5134 2009). al., et actin-rich (Rajan the midbody ring represents the Purple shape above cells. yellow located epidermal The structure Epid., ring. right. contractile the the to to represents is posterior Anterior and Scribble. Neurexin left Scrib: Nrx-IV: junctions; the larvae; tight giant with associated (2) protein Dlg: Lethal PATJ: Crumbs; IV; Lgl: Crb: E-cadherin; basolateral Coracle; E-cad: the Cora: large; at C; Discs kinase and/or protein junction atypical be adherens aPKC: might formed interface. cortex. Delta newly lateral and the posterior Notch at the between initiated at interaction apical enriched cytokinesis, the are During whereas aPKC (B) cortex, and lateral Par6 anterior Par3, marker from the markers junction switches at septate which enriched the of becomes mitosis; polarity Dlg during cell the antero-posterior the to SOP, to localize dividing apical-basal determinants the basal-to- fate of promotes cell cortex which Both anterior Neuralized, Delta. by of and transcytosis red regulated Numb apical partially the inhibitor is (containing Notch signalling cell the Notch pIIb by of blue the manner the in unidirectional (containing present The cell Delta nucleus). pIIa ligand the the in by Notch nucleus) of activation differential the organ sensory of interface Drosophila formed cells. newly daughter the precursor at Signalling 4. Fig. ou iieaymtial,adcl aeaqiiinrle on relies acquisition fate cell and asymmetrically, divide notum A esr ra rcros(Os fthe of (SOPs) precursors organ Sensory (A) nefc,aoetemdoy ewe h Iacl n pIIb and apical cell the pIIa the at 2012). between transiently al., midbody, et the resides (Couturier above Notch interface, cytokinesis cytokinesis, at Notch During initiated that revealed is construct signalling (GFP)–Notch newly protein the will fluorescent cytokinesis at we signalling during below, interface. Delta–Notch discussion cell cell the the daughter formed on In pIIa only 2009). the concentrate al., which et into (Sara), (Coumailleau and activation partitions receptor Notch for 2013; Furthermore, endosomes, unequally anchor multivesicular 1994). al., Smad of 1996; al., subset with al., et a marked et et into Rhyu internalized (Cotton Frise are 2012; 1996; Delta mitosis al., al., et unequally et during Couturier is Guo 2013; turn, cell al., and in et cell which, this Couturier pIIa Numb, by the by Notch in cell inherited pIIb activated receptor the is in transmembrane Notch inhibited Delta; the divide ligand its the of through on SOPs relies activation acquisition Schweisguth, fate an and Cell differential (Gho 1996). 1990). produce cell al., pIIa et to Schweisguth posterior 1998; a axis Posakony, and cell antero-posterior pathway pIIb anterior the and along signalling asymmetrically Hartenstein in (SOP) Notch precursor 1989; organ the sensory the Drosophila of of division cell following activation differential eut nNthatvto.Ftr tde ie tanalyzing will SOP at the aimed of cytokinesis studies during Future dynamics polarity activation. hence membrane and Notch Notch, in and/or pIIa– Delta polarity results formed with newly of interacts which the distribution interface along pIIb apical-basal polarity, apical-basal an i.e. of markers, (Bellaı possibility the 4A) the at (Fig. raises enriched and cortex becoming cortex lateral lateral Par3/Bazooka marker posterior anterior determinant junction the apical septate at the enriched the becoming with apical- large mitosis, the Discs of of during atypically remodelling segregation polarity the partial is unequal basal SOP-specific above the a polarity interface requires Indeed, Numb SOP apical mitosis. the that Interestingly, during al., at possibility remodelled et 4B). the activated (Fig. (Benhra raise be function data midbody also of These can 2013). gain and al., Notch Notch AP-1, et complex in Cotton signalling adaptor results 2011; Notch clathrin AP-1 the for of of platform loss that depleted SOPs a in been there as have accumulates 2010). have Notch act al., because side to cytokinesis apical et during proposed the (Benhra at been junctions cells also adherens basal-to-apical epithelial new in the the (Le Furthermore, cytokinesis Delta promotes at of 2003), cell transcytosis pIIb Neuralized, Schweisguth, the determinant and by fate Borgne inherited cell unequally The is 2009). is which al., that in et activation midbody (Rajan Notch the and SOP an of trafficking level of Delta the formation open for at required the an and control remains above Wasp interface 2012) structure actin-rich and al., basolateral Arp2/3 et the Indeed, Couturier at question. 2013; whether al., initiated However, et 4B). solely (Fig. (Couturier is choice order signalling fate in cell Notch asymmetry interface an binary pIIa–pIIb 2012). creates basal regulate Numb al., the to which along et in localization model (Couturier Notch a in cells Notch to led daughter Notch of data symmetric SOP These distribution hence both symmetric and in pIIa. interface, the activation in pIIa–pIIb in basal activated pIIb the results exclusively along the Numb is of of Notch of membrane endocytosis consequence, Loss plasma a the the As regulating from cell. Notch by removes basal Numb, the Notch, to delivered furrow. is endocytosed been cytokinetic has that Notch and cell, eeteeatwr htmd s fafntoa green functional a of use made that work elegant Recent ora fCl cec 21)17 1753 doi:10.1242/jcs.151472 5127–5137 127, (2014) Science Cell of Journal ua ou Fg )(atnti n Posakony, and (Hartenstein 4) (Fig. notum pupal cee l,20) This 2001). al., et ¨che Drosophila

Journal of Cell Science are spaeo ih ucin)fo ohrt agtrcells, exciting daughter Another integrity. to tissue the of mother maintenance diffusion from the ensures the junctions) investigating junction which of tight transmission or in adherens the (septate and barrier resides polarity of cell of challenge remodelling etc.) Another degradation, components. protein trafficking, ie ubro eldvsosadt retteei re to and order size in appropriate these with orient tissue to a shape. mitogenic and repair) divisions (or given cell generate undergo a ultimately of to order number decode in given source, and the a from sense distance at can sometimes the signal, cells in epithelial junctions important. how be tight Further also will and/or apically. abscission and septate midbody occur occurs the of of they maturation abscission role vertebrates, the and in investigating junctions) maturation whereas because adherens midbody informative, basally, very to invertebrates, be also apical in will junctions cytokinesis cell (tight epithelial vertebrate junctions) adherens to is and basal are it that as junctions between (septate comparison forces, invertebrate A regulates 2013). tensile negatively of al., bridge to et intercellular (Lafaurie-Janvore dissection subjected cytokinesis the in is molecular tension that that the tissue known with a proceeding in abscission in lies challenge ucin n sascae ihteaqiiino a of acquisition the with adherens spot-like associated to is adjacent adherens and the apical, to than basal junctions just rather of place cilia role takes junctions neuroepithelium, the mitosis mouse to after In regards re-synthesis junctions. with adherens acquisition nascent fate the cell the and of cytokinesis polarity and signalling. geometry Delta–Notch the on interface of adhesive impact new the decipher to help COMMENTARY b,M,Mkn,A,Hli-asd,F,Kmj,K,On-wsia Y., Ohno-Iwashita, K., Kamijo, F., Hullin-Matsuda, A., Makino, M., Abe, References le contre from Ligue ApiNotch). funding La SVSE2 core (CNRS), by Scientifique Labellise funded Recherche (Equipe is la Cancer R.L.B. de S.L.B.; National funds Centre 1 the Rennes of University The Funding interests. competing no declare authors The interests comments. and Competing reading critical for France) Bordeaux, (NeuroCentre 862, Montcouquiol du U M. Biologie INSERM and de Magendie France) (Institut Genetics Marseille, Lecuit of T. de Institute France), the Developpement Rennes, of (all of Tassan J.-P. Development Pacquelet, and A. Chesneau, L. thank We Acknowledgements Ca and the forces) decipher following (e.g. (pulling junctions to mechanics help the adherens the will biochemistry of and contacts during junctions contributions adhesive cells respective operate new adherens of neighbouring that formation of from forces measure mechanism disengagement to tensile able and Being question. the cellular open timing an the remains at disengagement the First our arise. as However, challenges level, cells. new epithelial increases, in recent process knowledge cytokinesis, this of of mechanics the dissection genetic in studies the confocal with time-lapse quantitative microscopy and non-invasive combining By remarks Concluding (Wilsch-Bra fate cell 2012). al., progenitor basal delaminating aaa . iuo . iaai .adKbysi T. Kobayashi, and A. Miyawaki, H., Mizuno, K., Hanada, ial,aohrcalnei oudrtn,a h iselevel, tissue the at understand, to is challenge another Finally, nte xml lohglgt h nepa between interplay the highlights also example Another Drosophila e,adTeNtoa eerhAec AR (Blanc (ANR) Agency Research National The and ´e), aepvdtewyt h nesadn of understanding the to way the paved have 2+ dpnetahso,membrane adhesion, -dependent 21) oefor role A (2012). uigret ¨uninger otre,L,Vdvr .adShesuh F. Schweisguth, and N. Vodovar, L., Couturier, Fu F., Coumailleau, R. Borgne, Le and N. Benhra, M., Cotton, A. Guichet, and K. Legent, B., and Benoit, P. J., J. Jouette, S., Thiery, Claret, E., Perez, F., Pincet, O., Eder, A., W. M., Thomas, S., Fukuda, Y. I., Chu, Kouranti, M., Machicoane, D., Dambournet, L., Chesneau, rwl,E . afr,A . aru-oe,B,Tjahh .adEhr,A. Echard, and S. Tajbakhsh, B., Gayraud-Morel, L., A. Gaffuri, F., E. Crowell, F. Schweisguth, and K. Mazouni, L., Couturier, J. S. Doxsey, and B. W. Huttner, W., G. A. 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Resurrecting C. in degradation Biol. midbody Cell cytokinetic J. promote regulators Apoptotic of godfather the to rider easy from (CPC): mitosis. complex release. passenger HIV-1 chromosomal and cytokinesis in USA ESCRT-III Sci. Acad. and Natl. Alix for requirements h tatcsud oiopae.I yolsi otniyadtissue and continuity Cytoplasmic I. pealei. Loligo squid, differentiation. Atlantic the machinery. ESCRT the for role a budding: mitosis. retroviral during size and shape cell of 29 function the Exploring (2014). separation. cell daughter to road the ogln .L,Gri-ays .adEgr,U S. localization. U. and Eggert, composition and lipid S. their regulate Garcia-Manyes, L., M. Coughlin, asymmetry. cell epithelial of generation the gametes. female from insights new positioning: cells. follicle Drosophila syncytial 4086. in movement protein 441. phosphatidylinositol- cytokinesis. of during furrow accumulation 1396-1407. cleavage the the to in 4,5-bisphosphate domains lipid sphingomyelin-rich ahrnctnncmlxi ioi ai-ab aiekde pteilcells. epithelial kidney Chem. canine Biol. Madin-Darby J. mitotic in complex cadherin-catenin abscission. orderly ensure to midbody the to Sci. Cell J. abscission. cytokinetic control that mechanisms salsmn,mitnne n remodeling. and maintenance, establishment, 21) erlzdpooe aa oaia rnctsso et nepithelial in delta of transcytosis apical to basal is promotes and Neuralized homeostasis (2010). 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