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e ae,C 61,USA. 06519, CT Haven, New Ato o orsodne([email protected]) correspondence for *Author USA. 06511, CT Haven New , of e ae,C 62,USA. 06520, CT Haven, New o obcei oaiepoen otecl pole? cell the to Ge localize do How COMMENTARY ß 2 1 Losick, environments 2013). and Jacobs-Wagner, Rudner diverse and 2009; al., Campos to et of 2010; Shapiro aspects 2006; al., various et and for (Matsumoto critical and bacterial is in organization bacterial arrangements spatial , This subcellular DNA, cells. display of , components can cellular (proteins, metabolites) that appreciated kinds well now various can is – It functions sorted. cellular hence misconception be -enclosed – of intracellular which lacks in the This generally organized, that cells observation tiny from the cells. these from originated removed randomly eukaryotic naturally of far vessels compartmentalized simple , were bacteria distributed the that challenging ago, cells idea decades bacterial two antiquated than of more organization accumulate to spatial started elaborate the for Evidence Introduction Spatial localization, Polar organization cycle, cell Bacterial WORDS: KEY localization. of patterns spatiotemporal complex obtain self-organizing to bacterial cells the by adopted on strategies been regulation emphasis present have also an We that principles. with mechanisms literature, major the the in specific review described a we at (for Here, example, (for time pole). space cell and in cycle) patterns cell diffusion-and-capture the reproducible during on be example, dynamic, can secrets generate event so-called this their how to and reveal regulated occur a can to ‘recognition’ pole starting initial the in also how are there, Bacteria were that mechanism. complexes located protein through or poles proteins already the Thus, other to with recruited is interaction are appendages. their proteins . proteins cell localized specific bacterial differentiation, in polarly of question of Often, crucial a localization cellular is as growth regulated polar and achieved progression, the processes important and how cycle underlie constitute understanding – that cell cells virulence, regulation as rod-shaped cell cellular the of certain essential particular, for ends of In cell. the positioning platforms the precise – of domains the poles of subcellular is bags to in merely Central organization proteins are chemicals. they spatial and that their concept macromolecules organized, initial distributed highly the are randomly from cells far bacterial is that appreciated which well now is It ABSTRACT eateto oeua,Clua n eeomna ilg,Yl University, Yale Biology, Developmental and Cellular Molecular, of Department Universite Institute, Duve de 04 ulse yTeCmayo ilgssLd|Junlo elSine(04 2,1–9doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal | Ltd of Company The by Published 2014. rlieLaloux ´raldine 1 ´ n hitn Jacobs-Wagner Christine and 4 3 eateto irba ahgnss aeSchool Yale Pathogenesis, Microbial of Department ahlqed ovi,B10 rses Belgium. Brussels, B-1200 Louvain, de Catholique oadHge eia nttt,Yl University, Yale Institute, Medical Hughes Howard 2,3,4, * h eemnnso iiinst lcmn iC i MinJ via MinCD the placement 2012), site al., division and 2011) of al., et et determinants (Briley Santos cells the competent in (dos Maf in inhibitor ComN division programs example, regulator cellular For competence several events. in cycle cell subtilis involved important or is play pathways to other DivIVA tend different recruit proteins in hub themselves These roles can forth. so which and proteins, proteins, also multiple that for proteins. hubs mechanism non-polar general many of a the localization is determine subcellular Note the capture’ will state. explains steady and at B A ‘diffusion protein and of that of accumulation A polar concentration the dissociation partners of and The extent association interacting pole. of rates the the the and cell between at poles is the the pool at in B localized A protein pool the protein diffusing and the interactions, body the between protein–protein of exchanged Because affinity. of continuously a binding encounters a nature has it scenario, it until transient this which protein space for In or cytoplasmic B 1A). protein the protein (Fig. polar in a pole diffuses the with A at protein interaction present an to already recruited through simply complex are poles far cell so identified the proteins polar the interaction of protein–protein Most through variations capture and themes and poles: Diffusion cell the at localize proteins How u betv st eiwadilsrt h eea principles general the illustrate and review to polar is Instead, reviewed of 2013). objective Waldor, been number and our Davis have large 2011; 2009; Viollier, which the and Dworkin, Kirkpatrick of 2007; of Jacobs-Wagner, many survey and (Ebersbach far, a elsewhere so provide goal described Our to proteins 2011). not Viollier, important is and this regulation, Kirkpatrick of here 2011; of cycle adhesion al., number and et cell focus (Bowman large chemotaxis, including the virulence, bacteria, a differentiation, cell is in for processes which essential cellular . proteins, in is widely polar Commentary, vary specifically ‘polar of and as proteins to numerous of Localization (referred are poles subset hereafter) the a at proteins’ localize where that Proteins event division localizes. a each by constitute generated are – that cells rod-shaped of ends ehnsst rdc pcfcsaitmoa atrsof patterns spatiotemporal specific localization. produce protein be these to control to that strategies mechanisms starting possible bacterial examine during we only selected Then, been . are polar have several that time summarize mechanisms first control localization and we that Commentary, space this mechanisms In uncovered. in the central yet localization others The underlie bacteria, cycle; polar poles. localization in cell cell polar processes two the of cellular the of asymmetry of course and one location the dynamics only change at during accumulate to as reproducibly known such are time, proteins over some poles. Indeed, the at complex. proteins localize to cells bacterial by used eakby oepoen a ev splrlnmrsor landmarks polar as serve can proteins some Remarkably, ihntebceilctpam h elpls–terounded the – poles cell the cytoplasm, bacterial the Within h atrsdslydb rtisisd atra el a be can cells bacterial inside proteins by displayed patterns The hog h erimn fmlil rtis the proteins: multiple of recruitment the through Bacillus 11

Journal of Cell Science COMMENTARY oad h poiepl,floe yters ftesister the of rest segregated the is 12 by other followed the pole, while opposite pole ParB– the one duplicated towards the at of partition remains one The complexes replication, origin the 2010). Upon and origin, al., ParB. the chromosomal the et of of Schofield vicinity composed 2008; al., 2010; is et Ebersbach al., complex and 2008; et al., complex, et Bowman (Bowman partition and control segregation cycle chromosome in cell involved the are with that proteins similarity with recruits through pole structural interaction the at or origin an chromosomal sequence 2008; the anchors no also DivIVA, In shares Kearns, 2003). which Errington, et PopZ, and and (Ben-Yehuda Wu RacA 2003; protein the al., an DNA-binding Patrick through attaches the spore developing with DivIVA the interaction in 2008; Moreover, pole the proteins. at 2011). origin illustrated chromosomal the al., al., of et the reflect membrane Eswaramoorthy et not cytoplasmic do CM, and space; scale periplasmic (Bramkamp P, to membrane; not outer are OM, figures space; all of in extracellular rest in ProP E, the (e.g. schematics and protein depicted. the a poles is that of cell cardiolipin, Note case between as particular cytoplasm. The such and C, proteins. poles, membrane polar cytoplasmic the of the at localization occurri of for enriched composition cues assembly as in protein serve Differences higher-order in can (D) a envelope PopZ region. of (e.g. ro in formation the DivIVA a higher-order outside right: (e.g. of favored as and curvature representation such Middle negative Left: assemblies 2010). stronger curvature. protein Ramamurthi, of stronger large and in regions of (Huang HubP membrane regions (e.g. previously in membrane protein described preferentially in polar as a favored ( area), for are curvature affinity (gray assemblies of an cylinder protein through radius arrows) Higher-order the (double (B) showing transiently features. poles. cell poles polar cell the of the at trapped recognition at is the localized arrows) through single proteins by indicated polar (as of Localization 1. Fig. C A Diffusion andcapture Nucleoid occlusion and proteins) (chromosomal DNA Nucleoid ifsn rti Polarprotein Diffusing protein aggregating protein self-assembling or Oligomer ofa R parS n h togrngtv uvtr ( curvature negative stronger the and ) eune,lctdi the in located sequences, albce crescentus Caulobacter parS bnigprotein -binding D B Affinity forpolarfeaturesofthecellenvelope Negative curvature parS .crescentus C. C R C RC R , = 2/ = 1/ = 2/ C uvdarw)a h elpls(leaes oprdwt h ie fthe of sides the with compared areas) (blue poles cell the at arrows) curved , Sidewall .subtilis B. hog iest fmcaim,a ecie below. described as poles mechanisms, the can of at diversity proteins several bacteria position a have efficiently surprisingly, through to poles cell not them The of the perhaps cell. several exploit makes the and of what To features, rest examine directly? distinctive the must from poles different one the poles question, ‘recognize’ this to address proteins position landmark 2012). al., these that et (Yamaichi pole the different at machineries three these of elements localizing by and synthesis chemotaxis segregation, chromosome between link oiinn ftedvso iena h icl,weethe where Shapiro, midcell, and correct the In (Thanbichler 2008). the near al., lowest et Ebersbach for the site 2006; is essential division concentration is the inhibitor both segregation of at complete positioning fastening after tight PopZ-dependent poles their inhibitor, ParB– division both Because chromosome. u hti h liaepstoa nomto htallows that information positional ultimate the is what But rpoenageae eg ifle rtisin proteins misfolded (e.g. aggregates protein or ) A rti eg aA in ParA1 (e.g. protein A (A) OM CM E C P .Arw niaetefe ifso foioes C omto of Formation (C) oligomers. of diffusion free the indicate Arrows ). ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal Pole .coli E. OM CM E C P htpeeetal id noi anionic binds preferentially that ) .cholerae V. Key irocholerae Vibrio at thepoles(e.g.cardiolipin) Anionic phospholipidenriched Peptidoglycan Peptidoglycan turnover Lipopolysaccharide cardiolipin Protein withaffinity for parS ifsn ntecytoplasm the in diffusing ) opee locrya carry also complexes .cholerae V. .coli E. uPpoie a provides HubP , senergetically is ) hti already is that ) d-shaped ng h cell the ;

Journal of Cell Science rgesvl eoaie otecl oe Rmmrh and (Ramamurthi poles DivIVA cell septa, In as the new 2009). form idea: to Losick, not this relocalizes do support and progressively blocked filamentous is, become artificially poles cells the with is that Experiments at septum. or strongest, a cells which of dividing the absence actively the DivIVA is in in that septum curvature idea the at the Harry negative primarily with 2003; initially consistent yet where Errington, is not and localizes This DivIVA are 2003). (Hamoen that Lewis, division spores septum, and cell outgrowing in in the poles engaged the at at concentrates accumulation dependent on fdvso siiitd(aautie l,2009; al., et of (Ramamurthi localization polar the initiated the in in Whereas new is 2011). DivIVA regions a al., when curved division et position most Eswaramoorthy septal of a the to time, round redistributes that DivIVA mainly division then at division, the cell), are, from cell generated (which newly After poles site self- al., 2009). the et at the (Lenarcic al., localized cell remains et by the of Ramamurthi regions provided 1B). concave 2009; (Fig. most is the cell in mechanism the of localizes of DivIVA this attractive membrane rest protein concave the of assembling An to more illustration relative at the poles 2010). An for accumulate the affinity preferentially of Ramamurthi, an curvature proteins through some and poles the that (Huang is hypothesis cell the Oiae l,21) ute ugsigta higher-order a of that septum localization suggesting curvature-mediated or the is further oligomers. pole for DivIVA tetramer the 2010), required is DivIVA at al., study assembly the a curvature et structural membrane of with recent (Oliva the curvature A from intrinsic interactions 2009). distinct the al., stabilizing that et showed of (Lenarcic membrane possibility the of model to clustering owing ‘molecular-bridging’ regions the more-curved enhanced these A at that favored and is argues 2009). oligomers simulations 2010), DivIVA al., Monte-Carlo al., et on based et Wang et Oliva 2004; (Stahlberg and structures notion, larger al., 2004; into assemble Huang this al., further can oligomers et with 2009; of Stahlberg Consistent al., sensing 2002; 1B). oligomerizes (Fig. et long-range DivIVA 2010) large (Lenarcic cooperative, into Ramamurthi, assemble curvature achieve to Ramamurthi, need membrane and to would (Huang protein be change structures to the curvature diameter scale? Instead, a pole any of 2010). the protein detect a to size relative to average the sense small the able too at that protein is shows monomer negligible model protein nanometer-sized mathematical is DivIVA a a that Indeed, that protein could difference a idea of how the curvature need But the supports without This anchor. concavity stronger 2000). senses lack al., directly that et species as (Edwards distant such very homologs in DivIVA expression heterologous are upon tips hyphal These 2013). al., et (Fla tips, Holmes formed surfaces hyphal 2008; curved al., emerging et negatively and Hempel sensing future by at probably localizes mainly homolog ln h yidia elbd (where body whereas cell total bacilli, cylinder), cylindrical The in example, the geometry. for along their shape: their is of poles because cell ( the curvature of characteristic curvature A enhanced of sensing Direct COMMENTARY .subtilis B. enovo de C ftecl neoei togra h poles the at stronger is envelope cell the of ) .subtilis B. iIAsotnosyacmltsa h poles the at accumulates spontaneously DivIVA ln h elwtotadvso event. division a without cell the along tetmcscoelicolor Streptomyces C qas2/ equals nvitro in eie otyfo t curvature- its from mostly derives shrci coli Escherichia R .subtilis B. and ttehmshrclplsof poles hemispherical the at nvivo in R hc preferentially which , yhe h DivIVA the hyphae, sterdu fthe of radius the is .subtilis B. (Muchova n iso yeast and C seult 1/ to equal is rh 2003; ¨rdh, ´ el in cells tal., et R ehns hti ae nPp seby(aoxadJacobs- and (Laloux assembly PopZ on based is that mechanism in only coli not its shown E. was to et This Bowman essential 2013). 2013; al., Jacobs-Wagner, is and multimerization (Laloux localization PopZ polar partners. PopZ Jacobs- known and Laloux though 2008; even al., 2013), et Wagner, divergent Ebersbach evolutionary 2008; the al., et in (Bowman polar a into 2010). al., et assemble al., Ebersbach et 2008; al., Bowman et 2008; (Bowman docking partners of multiple availability assemble for the further enhances sites presumably oligomers that matrix resulting a the into and itself in with PopZ protein hub called self-assembling process mechanism crescentus the localization a for proposed polar by first This poles was 1C). the (Fig. to occlusion sorted nucleoid passively is be protein can large the self-assembly Thus, clusters as effects. such protein volume-exclusion of polymers for because through bulky the nucleoid cue outside that structures favorable notion positional more the big energetically a on of This based as is (nucleoid). formation serve concept The DNA also by localization. characterized chromosomal protein can are of feature poles devoid distinguishing cell largely the geometry, being their from Apart occlusion nucleoid and Self-assembly l,20) h oa niheto adoii sdie yits by driven insertion is its cardiolipin favors thermodynamically of which species enrichment curvature, polar intrinsic et bacterial Bernal The 2004; several 2007). al., et cytoplasmic al., of Kawai the 2000; poles Dowhan, in and enriched For (Mileykovskaya the are 2003; specific surrounding that 1D). al., (Fig. 2012) membrane with et Weibel, (Mileykovskaya envelope preferentially and cardiolipin Renner interact as cell such proteins the phospholipids some of of and recognition possible example, protein composition features another on the relies Therefore, pole-specific is localization envelope. polar poles cell of cell mechanism their the of of maturation envelope trait cell unique the of Another elements pole-specific for Affinity self-assembly. protein this to of added regulation through be process can control spatial of and part temporal second Commentary, the this in described this as that However, spontaneous. Note 2013). is Emberly, Emberly,process a and obtain Saberi and 2013; to al., (Saberi sufficient et Coquel alone are 2010; entropy poles the by at pattern nucleoid polar-localization of absence multimerize to the ability intrinsic and its particle, and a further. of expand diffusion more passive nucleoid can the of they the where addition from poles cell the excluded the are by at other accumulate surface. they grow each their , aggregates on to misfolded patches amorphous stick hydrophobic to the of tend As exposure and the proteins cytoplasm to misfolded the simple: owing is in principle diffuse The freely 2013). al., et Coquel coli E. al., et (Ebersbach 2013). content Jacobs-Wagner, of and DNA Laloux low independent 2008; of preferential is its regions by PopZ in Jacobs-Wagner, determined be of self-assembly to and appears localization instead Laloux and polar a curvature, 2008; the of Thus, al., filaments 2013). et (Ebersbach the blocked along in example, crescentus for region, the segregated DNA-free at non-polar only between any not form in can also matrix but PopZ poles fide bona A 2013). Wagner, uprigti model, this Supporting in poles cell the at aggregate passively also proteins Misfolded uprigteie fasl-raiiglocalization self-organizing a of idea the supporting , sarsl fncei clso Wnlre l,2010; al., et (Winkler occlusion nucleoid of result a as Eesahe l,20) oZsotnosyinteracts spontaneously PopZ 2008). al., et (Ebersbach ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal uator mutant .coli E. nsilico in .coli E. .crescentus C. el nwihcl iiinis division cell which in cells iuain eosrt that demonstrate simulations ak oZhmlg rany or homologs PopZ lacks .crescentus C. oZi loal to able also is PopZ u loin also but , nvivo in .coli E. 13 C. C.

Journal of Cell Science ewe oa n o-oa etdgya ean obe to techniques remains advancing in peptidoglycan composition resides chemical challenge non-polar 2012; next is in The and evidence al., difference determined. the et polar a stage, Wirth this and between at 2008; correlative However, al., 2012). mostly et al., Radhakrishnan et (Lawler have mechanism Yamaichi proteins 2006; this few by could al., poles a the far, proteins et at So localize thereby some poles. to proposed and cell that been the features at conceivable peptidoglycan mainly is accumulate pole-specific It bind species. directly ‘active’ less these considered al., is in et peptidoglycan Pedro polar (de septation Therefore, cell 1997). following sidewall the to redirected htsrea oa oaiaincues. elements localization chemical polar key as the serve the identify that to lack required currently resolution which spatial analysis, composition peptidoglycan of sdet ietpoenlpditrcina h oe twudbe would It pole. the at the interaction with protein– direct this correlates a whether to unknown due is it is cell although cardiolipin, the of frequency content in manner The overall localization 2010). cardiolipin-dependent al., polar a et Romantsov of in 2007; poles 2008; al., curved et the al., (Romantsov to negatively et localize two Mukhopadhyay least highly MscS, At 2006; 2011). Weibel, al., in and Renner et (‘microdomains’) (Huang clusters as COMMENTARY onso iiin(aesiadTeit 06.Because 2006). Theriot, and few 14 of a (Rafelski pool after the pole division that older so the of ages, at pole trapped rounds wall cell ultimately cell the becomes The to as ActA poles. proposed down the is slows towards ActA turnover grows, peptidoglycan wall older lateral the and the accumulation follow protein As of growth. result wall a as cell length cell entire the along peptidoglycan along the spots with discrete associates (Garcı at then secreted membrane, is the bacterium ActA of the cell, entry upon that a Lacayo suggests into 1993; into model pathogen al., and prevailing et The cell Kocks 2012). host 1992; al., the et al., comet inside et a (Kocks bacterium in cells the resulting neighboring propels pole, that cell the tail the at induces actin ActA host cells, Gram-positive of mammalian polymerization inside the is slow pathogen of this ActA the When protein explain surface to pathogen the proposed of been This polarization has poles. cell mechanism the toward the guided multi-step along gradually constituents be wall could Accordingly, cylinder cell 2006). cell of with associate Theriot, rounds stably and that several (Rafelski proteins over division progressively peptidoglycan of and is of insertion growth material lateral blocks’ continuous wall ‘building through cell new poles the the towards It that a 2A). pushed proposed (Fig. as poles been wall the toward peptidoglycan has proteins (non-polar) direct less-direct passively lateral alternative, to the ‘shuttle’ an use 1D), a as can (Fig. exploited localization strategy be protein can for envelope polar cue the of specificities sidewall Whereas the from transmission Growth-dependent of as of insertion such absence because the bacteria is some is bacteria in This rod-shaped material synthesis. most peptidoglycan peptidoglycan of envelope cell poles new the localize of the to particularity poles at another cell the Indeed, at there. wall proteins peptidoglycan the This of maturation accumulation level. local their single-cell for 2012). the poles Weibel, the exploit and the also (Renner at might at content and proteins lipid lipids other correlated particular unique: of not enrichments also is polar phenomenon are the whether proteins see to interesting atracudas xli h atclrcmoiinand composition particular the exploit also could Bacteria ´ -e otloe l,21)adeetal ped uniformly spreads eventually and 2011) al., et Portillo a-del itramonocytogenes Listeria Rflk n hro,2006). Theriot, and (Rafelski .coli E. rtis rPand ProP proteins, .coli E. is h oe(tihure l,1999). al., from et away protein (Steinhauer entire 1993; the pole the of the diffusion along al., lateral proteolysis preventing specific surface, at through unclear et cell IcsA achieved remains of is maintenance It (Goldberg the pole 2001). but the occurs, al., membrane positioning et initial and Charles this periplasm outer how 1999; the al., into the et Steinhauer its in before pole insertion one functional to the in targeted IcsA, ActA of Interestingly, actin- counterpart dynamics. the a growth such connect bacterium, pathogen of the temporally motility of rate might propelling powered growth a mechanism the of on formation localization depends the – hence, tail and actin – ActA of polarization oaie ttenwcl oe fe iiin(hnihe and (Thanbichler division most after poles The remains cell by partners, new 2B). interacting given the (Fig. with at In is along localized separation machinery. FtsZ, mechanism division homolog cell the this after of of components pole illustration localization new in straightforward result freshly a could pole cytokinesis before ‘new’ localized at stably site a protein division a and the Therefore, mother division. at of its site the invariably from at cell pole formed daughter ‘old’ each an divide, inherits bacteria rod-shaped site When division the from Inheritance el omdcl oe ntepoey(..Tp in TipN (e.g. progeny the the with associated in remain poles division cell cell formed before newly midcell in the and ActA at more (e.g. localized becomes aging stably and cell laterally during and inserted growth inert of is more cycles peptidoglycan through new proceeds as cell the division, as and poles growth the cell towards from directed resulting proteins polar division. of Localization 2. Fig. A rti soitdwt h etdgya sprogressively is peptidoglycan the with associated protein A (A) B A pole ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal pole Old Old Transmission fromthesidewall Inheritance fromthesiteofdivision Key .monocytogenes L. to lateralinsertionofnewmaterial Displacement ofpeptidoglycanowing Peptidoglycan-associated protein Protein localizedatthenewpole hglaflexneri Shigella New pole New pole New pole .monocytogenes L. New pole Growth and Division division per ob directly be to appears , nietehs othe to host the inside .crescentus C. .crescentus C. pole Old pole Old .()Proteins (B) ). ). the ,

Journal of Cell Science tZrn,adtn oacmlt tteplspsil ya by 2013). possibly al., et poles the (Chiu the of sensing trapping negative-curvature at through involve accumulate clusters could to that the process tend of diffusion-and-capture area and the excluding ring, these membrane, Instead, FtsZ the division. in future freely of diffuse sites clusters the or ring FtsZ cytokinetic in clusters spheroides membrane-associated o eeomna n elcceporm uha h polar the as such beacon programs a as cycle acts coiled-coil cell it where membrane-bound and pole new developmental the the at for cycle, present is cell TipN protein in the TipN of landmark polar beginning the to by pole crescentus illustrated localization new is C. their the This at progeny. couple localization the their then in directly ensuring can not thereby are cues, division that division cell Proteins 2011). in al., involved et Goley 2006; Shapiro, COMMENTARY opl silto steMnsse in system Min the pole- is of poles oscillation case cell well-studied to-pole two A 2011). the Søgaard-Andersen, between and (Lenz oscillate proteins some Remarkably, and binding switch membrane nucleotide through oscillations Pole-to-pole of clustering polar the drive division, at in inheritance this self-assembly two spontaneous and Thus, localization, of polar protein generations. their for several consequence keys to recurrent after leads division progressively retention a This future and 2007). As accumulation to al., confined et 2009). (Thiem appear al., sites al., clusters et Wang larger et 2008; organization, Sourjik, pre-existing and Greenfield from Thiem 2007; away of 2008; al., formation far et the favored and (Thiem entropically ones, ones old is expand clusters to or new clusters new build manner to concentration-dependent a Armitage, in chemoreceptors self-assemble and spontaneous: stochastically (Sourjik is for arrangement body are periodic cell This are 2010). clusters few the case a along but Chemoreceptor positioned 1993), regularly Shapiro, chemotaxis. the and crucial (Maddock in is polar that predominantly function be membrane their the a at for display to cells, organization division- bacterial spatial in remarkable structures appears any signaling largest the of among This in independently chemoreceptors factor. spontaneously, associated occur a inherits can cell daughter pole. each newborn a its way, at is 2010), This marker 2007). that al., TipN cytokinesis al., et bacteria during et Yeh Gram-negative (Gerding membrane 2006; in outer complex al., for divisome Tol–Pal et responsible the the Lam is of 2006; requires step component al., and This et division midcell. (Huitema the cell at on cell-size- relocalizes a dependent predivisional in and the late possibly pole, manner, In new 2010). of dependent the al., from et delocalizes segregation TipN Schofield cells, 2010; unidirectional al., et the (Ptacin ParB– promotes 2006). al., duplicated et also Lam 2006; al., et TipN (Huitema flagellum a of biogenesis vrg,telws ttemdel loigdvso tthat at division time the that allowing on reveal midcell, studies is, modeling the inhibitor and Experimental at division only. location lowest MinC the the division average, of oscillations, the pole-to-pole concentration along continuous the carries of proteins MinD Because and of MinC. pool poles inhibitor a between system, this oscillates In segregation rapidly 2013). chromosome al., (Raskin facilitate et site might Ventura division and (Di 1999) the Boer, of de positioning and proper the to contributes navraino hsmcaim h icl oaiainstep localization midcell the mechanism, this of variation a In a eetybe hw ob needn fthe of independent be to shown been recently has Hieae l,20;Lme l,20) tthe At 2006). al., et Lam 2006; al., et (Huitema parS .coli E. .coli E. atto ope oadtenwpole new the toward complex partition raso hmrcpos hc are which chemoreceptors, of Arrays . neetnl,telclzto fthe of localization the Interestingly, . .coli E. Fg ) which 3), (Fig. Rhodobacter i.3 oet-oeoclaintruhncetd wthand switch nucleotide binding. through membrane oscillation Pole-to-pole 3. Fig. salsmn fteMn oa a ol efclttdb a by facilitated be could cap polar MinD the layers waves of 2012). spontaneous lipid establishment form Frey, artificial proteins and Min Halatek on Ventura idea, 2011; Di this al., 2008; with et al., Consistent Loose et collective 2011; Varma Sourjik, 2005; of and Kruse, and cycles Howard to 2001; sufficient and obtain be oscillations geometry might pole-to-pole membrane to cell explain the the at needed interplay Thus, MinD–MinE unbinding. the is and that binding than feature suggesting membrane away 1999), polar Boer, further de striped are no and a adopts (Raskin poles system re- pattern Min cell the localization pole? cells), preferentially When filamentous can in effect 2002). the (e.g. but MinE-destabilizing normal MinD al., at the membrane et from that zone away (Corbin the indicate furthest What MinD the on cells re-assembled. assembles new round anywhere has a mutant MinD localize of where in formation cell Experiments the and the the on dissociates of triggers cycle new pole, side a starting the before other cytoplasm reaches the et in eventually Park freely 2011; diffuses MinE al., in and et on, 2012). (Park with so model and jungle’ al., interacts the membrane, of the ‘Tarzan then at so-called dimer a MinE and MinD next (Hu cytoplasm. the the hydrolysis destabilizes the from ATP monomers into stimulates MinD at membrane and releasing MinD MinD thereby al., zone to et 2001), binds that Hu Lutkenhaus, protein ATP-bound 2002; of MinE al., from The edge et extends 2003). (Hu of the al., cell that et the zone of Lackner Dimers center 2003; a the in towards 2013). pole membrane the the Zheng, bind cooperatively and process self-organizing multi-step a (Shih is Min of behavior oscillatory hnarw) iEte id otenx iDAP trtoso this of Iterations MinD-ATP. next the 1, to (Step binds ATP then the triggers MinE in and arrows). monomers zone thin MinD-ADP dimer releases MinD-ATP which membrane-bound hydrolysis, a of edge soitswt h deo h e iDAPdmrzn tthe at zone dimer it 4). MinD-ATP until and new diffuses 3 cytoplasm, the the of (Steps pole into MinD-ATP edge membrane opposite released membrane-associated the is the of with MinE at out’ associates pole, i.e. ‘running one ring, After at MinE 3). dimers the and from 2 far reassociate (Steps nucleotide membrane collectively ADP/ATP at the dimers Following zone with and dimer arrow). redimerizes MinD-ATP thick MinD-ATP the 2, exchange, of (Step retraction membrane progressive the a to lead process 4 3 2 1 ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal In Time (~1–2minutes/cycle) .coli E. nvitro in iEbnsMn-T iesa the at dimers MinD-ATP binds MinE , nvivo in Loee l,20) The 2008). al., et (Loose Key Mihrtadd Boer, de and (Meinhardt MinD-ATP dimers) MinE (disassembles (monomer) Cytosolic MinD-ADP MinD-ATP (dimer) Membrane-bound 15

Journal of Cell Science oa oaiaini fe eprlyadsailyregulated, spatially and temporally often is time and localization space in Polar localization polar of Control under still 2012). is Frey, cues spatial and such (Halatek of Drew investigation Weibel, importance and 2003; the Renner although 2007; al., Marshall, 2012), which and et Cytrynbaum to 2005; (Mileykovskaya al., cardiolipin, et binds as preferentially such MinD protein polar-nucleation specific COMMENTARY ih et aiiaeo rvn rti sebya given at assembly 16 protein prevent or on times. facilitate occlusion and example, patterns to locations polar nucleoid modulating be for for or strategy might general 1B) a based, (Fig. Hence, 1C). localization sensing (Fig. curvature polar self-organizing or membrane for complexes process of key large recurrent mechanisms into a as assembly emerges protein structures above, assembly described protein of As modulation simply spatial and can Temporal the or in itself ‘encoded’ division. localization cell be polar from derive can drives that patterns for case (i.e. mechanism localization asymmetric in the of polar is propagation proteins the This unipolar) Hence, cells. 2010). misfolded al., daughter et the of stably the (Winkler to of are transmitted aggregates one be that will only large proteins cell of mother scenario, pole the describe stage another of old We later pole In a one matrix. at at cycle. 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Intrinsic or speculative. topic are time others hot in whereas examples, localization a strategies concrete protein regulatory have currently modulate several can is consider that we bacteria Below, in investigation. observed localization various to the environmental dynamics underlying to unipolar basis response from mechanistic in The or or proceeds signals. polar cycle change cell to or the diffuse as only bipolar) from pole et one (e.g. at (Shapiro localization accumulate development their proteins and Some 2009). bacterial al., for essential is which eldvso loihrnl otiue otetasiso of transmission the to contributes inherently also division Cell .crescentus C. .crescentus C. el eas t polar its because cells parS el,i invariably is cells, .coli E. atto complex partition Fg 4A) (Fig. rtoyi euneta vnulydgae PodJ degrades eventually that sequence proteolytic 02 hne l,20) oee,tepeiemechanism poles the precise at the localize determined. differentially However, be PodJ to al., 2006). of remains et forms al., (Viollier both PodJ et whereby of Chen function hence 2002; and localization proper oiyisoioeiainsae nfast-growing could In DivIVA tuberculosis state. of oligomerization phosphorylation its branching Hence, wall modify hyphal 2012). hyphal and cell al., the growth et of apical at (Hempel on foci arrest effects DivIVA dramatic upon AfsK, of with disassembly or tips kinase the For artificially to protein leads activated bacteria. synthesis, Ser/Thr be by eukaryotic-like can exploited which the be by can DivIVA in otherwise strategy studies Recent an this example, 4B). regulate (Fig. that GTPase- temporal to time suggest way under in phosphatases, localization a is polar provide kinases, spontaneous others, would cognate among this regulation, (GAPs), the proteins thereby of presence activating multimerize, the activity If to accumulation. proteins protein– or polar some spontaneous also through their of could poles impacting modifications ability the similar the to 1A), influence recruited Viollier, (Fig. point and interactions proteins some concern protein Kirkpatrick at date to 2011; are known that al., examples most et Although motility 2012). (Bulyha polarized in and adhesion involved transduction, and are signal regulation, that cycle proteins cell several complex of the control distribution of binding, intracellular changes nucleotide as or such state modifications, phosphorylation post-translational of variety modifications A post-translational through self-assembly Affecting ehrn ftesgeae ParB– (when segregated time right the the at and of PopZ pole) (new tethering diffusing pole matrix right promotes of the turn, A at in formation accumulation 2013). oligomers, PopZ Jacobs-Wagner, local of the and concentration (Laloux at higher a PopZ location ParA that in of at with accumulation oligomers results our the interacts 2013), pole that Jacobs-Wagner, ParA new proposes and 4C) Laloux Because (Fig. 2010; model 2010). al., et al., (Schofield et Jacobs- ParB– Shebelut and the (Laloux of the segregation pole the of parS during new accumulation occurs unipolar-to- the which the 2013), this at Wagner, to new that ParA linked the protein suggests directly at partitioning work is forms transition our matrix bipolar from PopZ the second Evidence at a pole. matrix chromosome-tethering that its PopZ requires but 4A), a function (Fig. inherit division through cells pole old daughter above, Jacobs-Wagner, and mentioned and Laloux 2013; al., event 2013; et 2013), cycle Ditkowski al., 2010; cell et al., particular et Fernandez-Fernandez a Iniesta 2010; to al., proteins coupling et polar (Bowman through of occur localization also the of can control spatial event and cycle Temporal cell a to Coupling a as from time, PodJ and of Conversion space PodJ. (PodJ in beacon long polar localization the polar for proposed modulate to strategy polar polar its 2010). would of Phosphorylation al., turn, et thereby synthesis in (Jani localization, Wag31. Wag31-dependent Polar and 2010). promote al., of oligomerization et (Jani localization phosphorylation an Wag31 in result might stimulates in which 2005), al., et increase (Kang enhanced is Wag31 rti laaeb pcfcpoessmgtas ersn a represent also might proteases specific by Protein atto ope Pai ta. 00 coil ta. 2010; al., et Schofield 2010; al., et (Ptacin complex partition L ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal oasotrfr (PodJ form shorter a to ) el,epeso fknssfrteDvV homolog DivIVA the for kinases of expression cells, .crescentus C. .coelicolor S. h nrae hshrlto of phosphorylation increased the , oZpoie neape As example. an provides PopZ parS S yacell-cycle-regulated a by ) ope sneeded). is complex Mycobacterium S ensures

Journal of Cell Science lgmrlast h omto falresrcuea h oe This pole. multimerization-dependent the a at to structure localization. regulation large polar or temporal a protein and of a spatial formation of provides the self-assembly the to the with leads 2, interaction oligomer diffusing Step to the In owing of protein. locally Concentration asymmetric increases event. (red) cycle oligomer cell protein a to segregation inherent DNA distribution during ParA (e.g. whose in asymmetric partner is a distribution with subcellular interaction protein–protein through in locally oligomer PopZ of (e.g. concentration oligomer The crescentus or (C) protein step. self-assembling hypothetical a phase a exponential indicates in mark Wag31 question of kinases specific the a in of upon expression phosphorylation (e.g. as signal such modifications by influenced COMMENTARY blt fsm rtist efasml n hrb olclz tthe at in localize homolog to DivIVA thereby The the and (B) Wag31, self-assemble synthesis. (e.g. polar to protein poles a proteins new acquire some after not of example did ability for that cell), progeny (top in focus appear Eventually, can aggregate. progeny accretions polar the polar existing in the produced onto proteins accumulate misfolded aggregates, in protein aggregates of old-pole protein to (e.g. bipolar accumulation Left: old-pole in event. PopZ division (e.g. cell localization a by regulation produced temporal naturally and localization. spatial polar for of strategies Possible 4. Fig. C B A .crescentus C. tuberculosis M. Asymmetry generatedbycelldivision Post-translational modifications Coupling withacell-cycle-associatedasymmetry Self-assembling protein oligomer Difffusing ,adteeyispoest omlieie a emodified be can multimerize, to propensity its thereby and ), .I tp1 h rti bu)hsa asymmetric an has (blue) protein the 1, Step In ). amorphous aggregates multimeric structuresor Proteins formingstable a nraeWg1popoyain.The phosphorylation). Wag31 increase may A smercplrpten a be can patterns polar Asymmetric (A) Step 1 Step 2 Step Step 1 New pole New pole Old pole Old pole .crescentus C. Signal ? .Rgt rpgto fan of propagation Right: ). .tuberculosis M. Old pole .coli E. .I h case the In ). modification Post-translational distribution asymmetric Protein with ol be could ) enovo de C. Old pole rmbt xeietlssadtersst nesadthe anticipate understand also to We effort mechanism. concerted theorists a localization take and each will spontaneous of it or experimentalists intricacies years, has otherwise few both bring progress events past from to the to exciting in Although cycle order made regulation 4). been in (Fig. cell temporal processes exploited localization and be specific can spatial to responses and dynamics physiological inherent Molecular 1–3). (Figs protein poles asymmetries drive cell to the principles polar at mechanistic localization of diversity a regulated use Bacteria spatiotemporally perspectives and Conclusion a good such be could (Ku accumulation polymerization, for spontaneous concentration a cytoplasmic candidates allow low not whose and might cofactor self-assemble any which self-assemble Bactofilins, without to poles. specific the propensity a at time their localize in at and modulating concentration pole their thereby modulate specific space, to a and cycle cell at the present during is time that factor interacting ParA-dependent a in recapitulated 2013). heterologous Wagner, be a could in localization fashion PopZ unipolar-to-bipolar the of model, pattern this with Consistent nivsiao fteHwr uhsMdclIsiue eoie nPCfor PMC in Deposited Institute. months. Medical 12 is Hughes after the C.J.-W Howard release from Belgium. the fellowship of of a investigator Health of French-speaking an of recipient the Institutes the of National was Department the G.L. Research by GM065835]. funded RO1 is number lab [grant Jacobs-Wagner the in Work Funding interests. competing no declare authors The interests Competing owing cited be of not reading could critical work for whose limitations. lab authors space all Jacobs-Wagner to to the apologize of We members manuscript. the the to grateful are We Acknowledgements 2012). Weibel, and Renner 2011; Weibel, composition and or shape (Renner variable with cells minimal of reconstruction novel from will benefit field The also inspections. visual to manual 2011; inherent biases precludes al., or unconscious automation the their et observations Importantly, 2012). Sliusarenko protein al., simple et 2008; Teeffelen from van al., quantify inferred et (Guberman levels be to single-cell measurements cannot and tools any population that both – essential of at – software part becoming dynamics post-imaging localization integral recent are respect, an that packages In become study. should localization of quantification patterns the Margolin, addition, 2012; subcellular In being al., 2012). Jensen, et protein or and Landgraf Swulius the 2010; 2012; misfolding on a al., overexpression, depending et remain tags, (Winkler upon visualized fluorescent – arise some artefactual accumulations to can fusion Currently, polar they as often future. concern – the patterns in localization expression distributions adjustable of subcellular or observations relevant physiologically near-native more ensure with should levels, combined et 2011), Charbon 1998; al., al., et (Griffin bacteria in visualization protein dyes for of organic of membrane-permeable development and The list tags cells. fluorescent less-bulky bacterial live the inside proteins of as tracking discovery increasing. keeps await proteins polar mechanisms identified additional that te efasmln rtismgtsmlryrl nan on rely similarly might proteins self-assembling Other seta oteivsiaino rti oaiaini the is localization protein of investigation the to Essential ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal h ta. 2010). al., et ¨hn nvitro in .coli E. and ytm(aoxadJacobs- and (Laloux system nvivo in sasbsdo the on based assays nvitro in 17

Journal of Cell Science oul .S,Jcb .P,Pie,M,Dmrz . iicl,M,Jlu T., Julou, P. M., Dimiccoli, J. A., Armitage, Demarez, M., and Primet, J.-P., C. Jacob, M. A.-S., Coquel, Leake, J., A. M. Roberts, S.-W., Chiu, hn .C,Hte,A . cdm,H . crt,P . ilir .H and H. P. Viollier, T., P. McGrath, H., H. McAdams, K., A. Hottes, C., J. Chen, hre,M,Pe M., Charles, Schultz, A., Løbner-Olesen, J., Wang, A., K. Scott, E., Brustad, G., Charbon, saaorh,P,Eb .L,Geoy .A,Slemn . olao K., Pogliano, J., Silverman, A., J. Gregory, L., M. Erb, P., Eswaramoorthy, aps .adJcb-anr C. Jacobs-Wagner, and M. Campos, dad,D . hmie,H .adErntn J. Errington, and B. H. Thomaides, H., D. Edwards, uya . o,E,Hnly .adSgadAdre,L. Søgaard-Andersen, and S. Huntley, E., Hot, I., Bulyha, D. Dubnau, and J. Hahn, J., M. and Dias, P., A. Prepiak, Jr, R. K., Briley, Daniel, C., Donovan, L., Weston, R., Emmins, M., Bramkamp, brbc,G,Bigl . esn .J n aosWge,C. Jacobs-Wagner, and J. G. Jensen, A., Briegel, G., Ebersbach, C. Jacobs-Wagner, and G. Ebersbach, omn .R,Prz .M,Pai,J . goao . ot-tgiw E., Folta-Stogniew, E., Ighodaro, L., J. Ptacin, M., A. Perez, R., G. Bowman, rw .A,Obr,M .adRtfed .I. J. L. Dworkin, Rothfield, and J. M. Osborn, A., D. Drew, J. F. Gueiros-Filho, and W. A. Bisson-Filho, T., V. Santos, dos K., Ginda, M., Bezulska, M., Donczew, J., Rydzak, N., Holmes, B., Ditkowski, omn .R,Lusuoa .I n hpr,L. Shapiro, and I. A. Lyuksyutova, R., G. Bowman, Y., Jones, S.-H., Hong, M., Fero, M., G. Gaietta, R., L. Comolli, R., G. Bowman, H., K. Downing, M., Koenig, M., Eckart, J., Zhu, R., L. Comolli, R., G. Bowman, ePdo .A,Qitl,J . Ho C., J. Quintela, A., K. M. Pedro, M. de Waldor, and M. B. Davis, L. D. B. Marshall, and N. E. Cytrynbaum, W. Margolin, and X.-C. Yu, D., B. Corbin, iVnua . nct . nra,H,Gdnz .J,Fish,M,Rh,K., Rohr, M., Fritsche, J., W. Godinez, H., Andreas, B., Knecht, B., Ventura, Di V. Sourjik, and B. Ventura, Di enl . Mun P., Bernal, R. Losick, and Z. D. Rudner, S., Ben-Yehuda, References COMMENTARY 18 grgto nEceihacl sgvre ydfuinadnucleoid and diffusion by governed H. is Berry, coli effect. and crowding Escherichia B. macromolecular in Rhodobacter A. Lindner, aggregation the L., through Moisan, FtsZ and proteins cycle. cell chemosensory sphaeroides of Positioning elccergltdprotease. cycle-regulated cell a L. Vibrio. Shapiro, and Enterobacteriacae in USA IcsA Sci. factor Acad. virulence Shigella of oaiainb sn eeial noe loecn mn acid. amino fluorescent encoded genetically E. a Chapman, using and ChemBioChem by C. localization Jacobs-Wagner, G., P. olao .adRmmrh,K S. K. Ramamurthi, and J. Pogliano, fgntcinformation. genetic of agtn fBclu utlscl iiinpoenDvV odvso ie in sites division to DivIVA protein division yeast. cell fission and subtilis coli Escherichia Bacillus of targeting atra elplrt n signalling. subtilis. and B. polarity of cell bacterial cells competent in division cell arrest Microbiol. Mol. to ComGA of MinCD. downstream inhibitor division the for action of mode new Microbiol. a and subtilis Bacillus J. Errington, efascaigpoenciia o hoooeatcmn,dvso,adpolar and division, caulobacter. attachment, in chromosome organization for critical protein self-associating polarity. bacterial of mechanisms temporal otiuet u-ellrlclzto fabceilscaffold. bacterial a of localization sub-cellular to contribute L. Shapiro, and R. L. Comolli, eset Biol. Perspect. placement. site division 102 oscillatory bacterial self-sustained in the involved generates system accurately that model depolymerization Bacillus of regulator posttranscriptional a ComN, subtilis. of localization Streptomyces polar in mediated segregation chromosome with growth coelicolor. the coordinates Scy, D. Zakrzewska-Czerwin Jakimowicz, P., Kedzierski, ur pn elBiol. Cell al. Opin. et Curr. H. H. pole McAdams, in changes H., function. sequential M. and dictating subdomain Ellisman, composition polar H., a forms K. PopZ Downing, Caulobacter H., J. Lee, L. pole. cell Shapiro, bacterial a and at 945-955. complex T. origin/ParB chromosomal Earnest, the anchors E., W. Moerner, yteEceihacl i system. Min coli Escherichia V. the Sourjik, and by W. D. Heermann, W., Nickel, division. cell bacterial in proteins localized coli. Escherichia in segregation rods. division. negative cell coli E. in dynamics MinDE explains coli. Escherichia round-shaped in 2008. system Min the of function odusrltdt rnpr functionality. transport sensitivity to increased related exhibits drugs mutant to synthesis cardiolipin putida Pseudomonas poles. cell the to anchors that 6114-6118. , .Bacteriol. J. 70 20) ellrplrt npoaytcorganisms. prokaryotic in polarity Cellular (2009). pnBiol. Open 20) yoiei inl rnaino h oJplrt atrby factor polarity PodJ the of truncation signals Cytokinesis (2006). zRjs . utd,A,Rms .L n eua A. Segura, and L. J. Ramos, A., Hurtado, J., oz-Rojas, ˜ rz . oi,J .adGlbr,M B. M. Goldberg, and H. J. Kobil, M., ´rez, 1556-1569. , 20) oe opnn ftedvso-ieslcinsse of system selection division-site the of component novel A (2008). ur pn Microbiol Opin. Curr. 81 1 12 98 a003368. , 21) yai nepa fPr ihteplrt protein, polarity the with ParA of interplay Dynamic (2013). 23-39. , 1818-1821. , 9871-9876. , 194 23 3 ur pn Microbiol. Opin. Curr. 130006-130006. , o.Microbiol. Mol. 71-77. , 3661-3669. , o.Microbiol. Mol. Cell 21) efognzdpriinn fdynamically of partitioning Self-organized (2011). 21) lgmrzto n ihrodrassembly higher-order and Oligomerization (2013). .Bacteriol. J. MOJ. EMBO 134 LSCmu.Biol. Comput. PLOS 21) salsigplriett ngram- in identity polar Establishing (2013). te .V n cwr,H. Schwarz, and V. J. ltje, ¨ MOJ. EMBO i press) (in . 21) ellrognzto ftetransfer the of organization Cellular (2013). o.Ss.Biol. Syst. Mol. ur pn Microbiol. Opin. Curr. 956-968. , 90 20) xlrto notesailand spatial the into Exploration (2007). 21) ellracietr mediates architecture Cellular (2011). 20) utsrne oye model polymer multistranded A (2007). 2-3.10.1111/mmi.12366 322-337. , nio.Microbiol. Environ. ´ 76 20) xlrn nrclua space: intracellular Exploring (2002). 25 o.Ss.Biol. Syst. Mol. k,J,Klmn .H and H. G. Kelemen, J., ska, rnsMicrobiol. Trends 173-189. , 377-386. , 179 19 16 21) hoooesegregation Chromosome (2013). Science 20) aA atra protein bacterial a RacA, (2003). 21) oaiaino protein of Localization (2013). 2719-2727. , 171-176. , 2823-2834. , ipy.J. Biophys. 20) oyei protein polymeric A (2008). 21) uclua protein Subcellular (2011). rc al cd c.USA Sci. Acad. Natl. Proc. 9 20) polymerization- A (2005). 686. , 21) atra polarity. Bacterial (2011). 299 9 e1003038. , 20) oa targeting Polar (2001). 20) Promiscuous (2000). o.Microbiol Mol. 532-536. , 14 7 21) Tae in (2011). MOJ. EMBO 457. , odSrn Harb. Spring Cold 93 9 15 726-733. , 1135-1145. , 21) a acts Maf (2011). 21) DivIVA- (2012). 1134-1150. , 101-108. , 19) Murein (1997). rc Natl. Proc. 20) A (2008). 21 20) A (2007). Cell . (2013). (2010). 1998- , 134 Mol. , enne-enne,C,Gos,K,Sujk .adClir J. Collier, and V. Sourjik, K., Grosse, C., Fernandez-Fernandez, epl .M,Wn,S-. ee,M,Gl .A n Fla and A. J. Gil, M., Letek, S.-B., Wang, M., A. Hempel, aon .W n rigo,J. Errington, and W. L. Hamoen, E. Frey, Z. and Gitai, J. and S. Halatek, N. Wingreen, J., Dworkin, A., Fay, M., J. Guberman, J. A. P. Boer, de and H. Niki, D., N. Pecora, Y., Ogata, A., M. Gerding, Garcı Fla oms .A,Wlhw . egt,R . hbsad . atn .A., K. Dalton, A., Thibessard, M., R. Leggett, J., Walshaw, A., N. Holmes, L., J. Parker, J., M. Naldrett, S.-B., Wang, V., Molle, S., Cantlay, M., A. Hempel, J. P. Lewis, and J. E. Harry, u . az .adLtehu,J. Lutkenhaus, and C. Saez, Z., Hu, J. Lutkenhaus, and P. E. Gogol, Z., Hu, J. Lutkenhaus, and Z. Hu, K. Kruse, and M. Howard, rfi,B . dm,S .adTin .Y. R. Tsien, and R. S., S. N. Adams, A., Wingreen, B. Griffin, E., G. Crooks, H., Shroff, L., A. McEvoy, H. D., H. McAdams, Greenfield, E., Abeliuk, J., M. Fero, S.-H., Hong, Y.-C., Yeh, D., E. Goley, Ba B., M. Goldberg, ikarc,C .adVole,P H. P. Viollier, and L. C. K. Matsumoto, Kirkpatrick, and H. Hara, Y., Sadaie, R., and Harashima, C. M., L. Shoda, F., Cantley, Kawai, C., C. J.-S., Dascher, Han, T., S. B., Park, M. W., D. Sahana, Abbott, K., C.-M., Kang, Hamasha, J., J. Lee, H., Eoh, C., Jani, L. Shapiro, and J. N. Hillson, A., Viollier, A. and Iniesta, K. S. Radhakrishnan, D., Matteson, S., Pritchard, E., S. Huitema, N. Wingreen, and R. Mukhopadhyay, C., K. S. Huang, K. Ramamurthi, and C. K. Huang, rh K. ¨rdh, iIAutatutr n euae i ciiyi ailssubtilis. Bacillus in activity Min regulates e00257-11. and DivIVA utlsi o ietydpneto tZo B 2B. PBP or FtsZ on dependent dynamics. directly not MinCDE-protein is subtilis robust of origin the Reports explain computational sequestration by revealed resolution. division sub-pixel bacterial at in analysis asymmetry image and noise PSICIC: of proteomics wall cell by revealed peptidoglycan to monocytogenes. Listeria ActA intracellular of Association Caulobacter A3(2). in coelicolor replisome Streptomyces the to HdaA of localization the crescentus. directs clamp sliding ilsi,M . emns .M,Gs,B n eee,G H. G. Kelemen, controlling and assembly multiprotein B. a of Gust, component M., key a A. is Scy Hemmings, protein Coiled-coil D., M. Gillespie, Fla and J. M. Buttner, Y.-G., new Jung, M., establish D. can Richards, and branching coelicolor. Streptomyces hyphal in growth for wall cell sites of zones mark DivIVA of Assemblies site. division the to proteins Min of recruitment apparatus-independent division for evidence subtilis: Bacillus of spores germinated hshlpdvsce euae yAPadMinE. and ATP 99 by regulated MinE vesicles by phospholipid ATPase its of stimulation requires phospholipid. MinD and of oscillation spatiotemporal dynamics. protein Min for models and mechanisms Streptomyces. in growth polarized eobnn rti oeue nielv cells. live inside protein recombinant microscopy. light super-resolution 7 with imaged J. network Liphardt, chemotaxis and E. Betzig, Microbiol. in L. Shapiro, involved and protein flexneri Shigella a IcsA, movement. of intracellular activity ATPase and E. localization in and constriction machinery during cell invagination the outer-membrane coli. of proper part for is required complex Tol-Pal trans-envelope The rti iaeAs euae oa rwhadhpa rnhn ntefilamentous the Streptomyces. in branching bacteria hyphal and growth polar regulates AfsK kinase protein 20) adoii oan nBclu utlsmrugmembranes. marburg subtilis Bacillus in Bacteriol. domains J. Cardiolipin shape. cell (2004). of regulation and identification Dev. PknB: and PknA N. kinases R. Husson, mycobacteria. in phosphorylation al. Wag31 et Microbiol. H. by BMC S. cycle Lee, peptidoglycan J.-W., cell Suh, polar J.-Y., Lee, Caulobacter S., Nyayapathy, initiation. for replication essential DNA is on 3893-3902. dependent which are activation, progression, kinase histidine Cell H. P. poles. bacterial to lipids of Biol. localization for mechanism mediated MinE. and curvy. MinD membranes of role coli: Escherichia in Bacteriol. membrane J. the to formation, Z-ring raelsadtersailregulation. spatial a006809. their and organelles e1000137. , 6761-6766. , adlPril,F,Clo . ’rzo .adPcirli .G. M. Pucciarelli, and V. D’Orazio, E., Calvo, F., Portillo, ´a-del 124 o.Microbiol. Mol. 2 20) atra it crpoen akftr lglu sebysite. assembly flagellum future mark proteins scar birth Bacterial (2006). e151. , 1025-1037. , 1 20) seta oeo iIAi oa rwhadmrhgnssin and growth polar in DivIVA of role Essential (2003). 741-752. , 80 ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal 186, 185 19 1680-1698. , 1692-1704. , 10 196-203. , 21) sebyo h albce eldvso machine. division cell Caulobacter the of Assembly (2011). 1475-1483. 20) h yoatru ueclssserine/threonine tuberculosis Mycobacterium The (2005). ru . ast .adSnoet,P J. P. Sansonetti, and C. Parsot, O., ˆrzu, o.Cell Mol. 327. , o.Microbiol. Mol. 63 1008-1025. , rc al cd c.USA Sci. Acad. Natl. Proc. .Bacteriol. J. . 20) oooia euaino eldvso nE coli. E. in division cell of regulation Topological (2001). 20) al agtn fMnpoen otecl oe in poles cell the to proteins Min of targeting Early (2003). 21) ihycnlzdMn rnfradMinE and transfer MinD canalized Highly (2012). 20) ellrognzto yself-organization: by organization Cellular (2005). 7 1337-1343. , 20) efognzto fteEceihacoli Escherichia the of Self-organization (2009). o.Microbiol. Mol. 20) oa agtn fDvV nBacillus in DivIVA of targeting Polar (2003). rc al cd c.USA Sci. Acad. Natl. Proc. 76 20) erimn fMn,a niio of inhibitor an MinC, of Recruitment (2003). 175 822-832. , .Bo.Chem. Biol. J. 21) armlclsta rfrtheir prefer that Macromolecules (2010). LSCmu.Biol. Comput. PLOS 21) oe pr:poaytcpolar prokaryotic apart: Poles (2011). 20) yai sebyo iDon MinD of assembly Dynamic (2002). 2189-2196. , odSrn ab eset Biol. Perspect. Harb. Spring Cold 19) pcfccvln aeigof labeling covalent Specific (1998). o.Microbiol. Mol. 49 21) oa eoeigand remodeling Polar (2010). Science 1523-1536. , 109 .Bacteriol. J. ¨rdh, K. rc al cd c.USA Sci. Acad. Natl. Proc. .Cl Biol. Cell J. 286 .Bacteriol. J. E2371-E2379. , 34675-34689. , 281 21) euainof Regulation (2010). 20) curvature- A (2006). 21) h Ser/Thr The (2012). 47 4 269-272. , .Bacteriol. J. e1000233. , rh K. ¨rdh, 110 19) Unipolar (1993). 190 ,37-48. LSComput. PLOS 168 21) The (2013). 185 E397-E406. , 7579-7583. , 533-536. , LSBiol. PLoS 693-697. , mBio (2013). (2008). (2007). (2008). (2011). 192 Mol. Cell b 2 3 - , , ,

Journal of Cell Science aauti .S n oik R. Losick, and S. K. Ramamurthi, A. J. Theriot, and M. S. Rafelski, tcn .L,Le .F,Gre,E . oo . cat . ool,L R., L. H. P. Comolli, Viollier, M., and M. Eckart, Thanbichler, E., K., Toro, S. C., Radhakrishnan, E. Garner, F., S. Lee, L., J. Ptacin, B. D. Kearns, and J. E. Lutkenhaus, J. and Patrick, S. Lovell, W., Wu, J. K.-T., Lutkenhaus, Park, and T. Holyoak, S., Lovell, P., K. Battaile, W., Wu, K.-T., Park, Veprintsev, A., T. Leonard, P., Dutow, M., S. Freund, S., Halbedel, A., M. Oliva, S. N. Wingreen, and C. K. Huang, R., Mukhopadhyay, Muchova Dowhan, and W. Margolin, D., B. Corbin, X., Fu, I., Fishov, E., Mileykovskaya, W. Dowhan, and E. Mileykovskaya, A. P. Boer, de and H. H. Meinhardt, Hara, and A. Nishibori, J., Kusaka, K., Matsumoto, W. Margolin, ikarc,C .adVole,P H. P. Viollier, and L. C. Kirkpatrick, COMMENTARY adc,J .adSaio L. Shapiro, and R. J. Maddock, P. Schwille, and K. Kruse, P. M., Schwille, Loose, and K. Kruse, J., Ries, E., Fischer-Friedrich, J., M., Errington, Loose, J., L. L. Søgaard-Andersen, Wu, and P. M., Lenz, Shaw, V. Y. L., Brun, Visser, and S., D. Halbedel, Klein, R., J., Lenarcic, A. Hinz, E., D. Larson, J. L., Paulsson, M. and Lawler, A. T. Baker, P., Chien, B., Okumus, D., Landgraf, C. Jacobs-Wagner, and B. W. Schofield, H., Lam, C. Jacobs-Wagner, J. and A. P. G. Boer, Laloux, de and M. D. Raskin, L., Soo, J., L. M. Lackner, Footer, A., M. Tsuchida, J., Zhu, G., A. P. Soneral, I., C. Lacayo, aauti .S,Lcyr . tn,H .adLsc,R. Losick, and A. H. Stone, S., Lecuyer, S., K. Ramamurthi, Ku P. Cossart, Cossart, and H. and Ohayon, P., H. Gounon, R., Ohayon, Hellio, C., P., Kocks, Berche, M., Tabouret, E., Gouin, C., Kocks, n . ree,A,Mo A., Briegel, J., hn, ¨ u o uclua oaiaino atra protein. bacterial a of 106 localization subcellular for cue ActA. protein surface monocytogenes nepa ewe elft eemnn n yoyehmlgdie the drives homolog lysozyme a crescentus. and Caulobacter of determinant cycle fate division cell asymmetric a between interplay L. Shapiro, segregation. and chromosome bacterial E. W. Moerner, subtilis. Bacillus in division cell of MinE. by ATPase MinD the in of activation changes asymmetric conformational MinD-dependent cytokinesis. regulate uses spatially to oscillator MinE Min The (2011). structure. Lo crystal DivIVA and by W. revealed L. binding Hamoen, B., D. separation. microphase curvature-mediated J. through cells bacterial in iiinpoenDivIVA. Bara protein division and J. A. Wilkinson, vivo. in and vitro in dye fluorescent W. cardiolipin-specific the using by orange. acridine coli 10-N-nonyl Escherichia in the domains of localization the and proteins Min site. of oscillations division pole-to-pole the for model membranes. bacterial 194 pta euaosfrbceilcl iiinsl-raieit ufc ae in waves surface into self-organize division cell vitro. bacterial for regulators Spatial bacteria. in PodJ membranes. factor curved negatively W. localization to L. Hamoen, polar and D. Marenduzzo, the of domains crescentus. functional Caulobacter of Dissection localization. protein native reveals division Methods cell at molecules cell. of bacterial Segregation a of polarity the perpetuating and 124 establishing for essential phospholipid multimerization. the cycle-coupled 841. and cell through proteins localization Min coli vitro. A. in Escherichia membrane J. between comet Theriot, actin interactions and on polarization A. ActA Mogilner, and Y., geometry cargo tails. Xia, of Y., influence orientation: Lu, S., F. ope nteEceihacl cell. coli Escherichia the in . min the from Caulobacter in synthase wall cell a of localization of crescentus. polar site mediating the proteins M. at Thanbichler, ActA and protein surface monocytogenes assembly. actin Listeria directional of distribution protein. surface a product, P. Rev. Microbiol. FEMS u o rti oaiaini bacterium. a in localization protein for cue 95 19) .mnctgnsidcdatnasml eursteat actA the requires assembly actin monocytogenes-induced L. (1992). 20) fet fpopoii opsto nMn-ebaeinteractions MinD-membrane on composition phospholipid of Effects (2003). 6369-6371. , 1011-1023. , 13541-13545. , 1034-1049. , o.Bo.Cell Biol. Mol. Science ,K,Kutejova K., ´, a.Rv Microbiol. Rev. Nat. 9 480-482. , 21) h rc ftg npoenlclzto studies. localization protein in tags of price The (2012). MOJ. EMBO rc al cd c.USA Sci. Acad. Natl. Proc. 320 789-792. , .Bacteriol. J. .Bo.Chem. Biol. J. 23 nu e.Biophys. Rev. Annu. ,E,Sot .J,Bangn .A,Lws .J., R. Lewis, A., J. Brannigan, J., D. Scott, E., ´, shl . an,J,Lsr . ik . esn .J. G. Jensen, S., Wick, K., Leser, J., Kahnt, E., rschel, ¨ 29 o.Microbiol. Mol. 614-629. , 36 21) atfln,auiutu ls fcytoskeletal of class ubiquitous a Bactofilins, (2010). Microbiology 327-339. , o.Microbiol. Mol. 193-205. , k I. ´k, Cell .Cl Sci. Cell J. .Bacteriol. J. 9 e J. we, ¨ 565-577. , 20) iJ(vD satplgcldeterminant topological a is (YvjD) MinJ (2008). 68 185 20) lgmrzto fteBclu subtilis Bacillus the of Oligomerization (2002). 20) atr omto nEceihacl:a coli: Escherichia in formation Pattern (2001). 19) oa oaino h chemoreceptor the of location Polar (1993). 20) ehns fplrzto fListeria of polarization of Mechanism (2006). MOJ. EMBO o.Microbiol. Mol. 521-531. , 21) eoigCuoatrdevelopment. Caulobacter Decoding (2012). 20) eaiemmrn uvtr sa as curvature membrane Negative (2009). 278 21) eprladsailoclain in oscillations spatial and Temporal (2011). 735-749. , a.Cl Biol. Cell Nat. Science 21) pnl-ieaprtsguides apparatus spindle-like A (2010). 148 20) oaiaino iIAb targeting by DivIVA of Localisation (2009). 21) ptoeprlcnrlo PopZ of control Spatiotemporal (2013). 61 21) rti efognzto:lessons self-organization: Protein (2011). o.Microbiol. Mol. Cell 22193-22198. , 105 21) etrsciia o membrane for critical Features (2010). 20) iulzto fphospholipid of Visualization (2000). 1110-1117. , 59 98 807-813. , 182 699-710. , Science 301-316. , 146 40 14202-14207. , 28 1172-1175. , MOJ. EMBO 259 315-336. , 2272-2282. , 396-407. , 1717-1723. , 70 12 o.Microbiol. Mol. 20) adakprotein landmark A (2006). 323 rc al cd c.USA Sci. Acad. Natl. Proc. 1166-1179. , 21) ehns fthe of Mechanism (2012). 791-798. , ee Dev. Genes 59 20) ii localization Lipid (2008). 20) ii oan in domains Lipid (2006). 20) ATP-dependent (2003). 29 1354-1357. , .Cl Biol. Cell J. 1262-1279. , 1988-2001. , 20) h dynamic The (2008). 20) Geometric (2009). 21) Choosing (2012). 19) Polarized (1993). 22 85 .Bacteriol. J. 212-225. , 271-281. , 201 Biophys. (2006). (2012). (2008). 827- , Nat. Cell a efee,S,Savt,J .adGti Z. Gitai, and W. J. Shaevitz, Teeffelen, S., van he,S,Knnr .adSujk V. Sourjik, and D. Kentner, S., Thiem, V. Sourjik, and S. Thiem, L. Shapiro, and M. Thanbichler, luaek,O,Hirt,J,Eoe,T n aosWge,C. Jacobs-Wagner, and T. Emonet, J., Heinritz, O., Sliusarenko, M. Zheng, and Y.-L. Shih, ori,V n riae .P. Kutejova H., J. Stahlberg, Armitage, and V. Sourjik, e,Y-. ool,L . onn,K . hpr,L n cdm,H H. H. McAdams, and L. Shapiro, H., K. Downing, R., L. Comolli, Y.-C., Yeh, ag .B,Cnly . odeg . ee,M,Gl .A n Fla and A. J. Gil, M., R. Letek, N., Mukhopadhyay, Nordberg, S., and Cantlay, S.-B., S. Wang, N. L. Wingreen, H., Shapiro, Wang, and N. Sternheim, H., P. Viollier, D. K. Young, and C. K. Huang, A., Varma, J. G. Jensen, and T. M. Swulius, B. M. Goldberg, and W. A. Varga, T., Pham, R., Agha, J., Steinhauer, hblt .W,Gbra,J . a efee,S,Yknn,A .adGitai, and A. A. Yakhnina, S., Teeffelen, van M., J. R. Guberman, W., Losick, C. and Shebelut, H. H. McAdams, L., Shapiro, aaci . rcnr . igar,S,Mo S., Ringgaard, R., Bruckner, Y., Yamaichi, J. Errington, and J. L. Wu, M., Fernandez-Suarez, M., S. Fortune, B., B. Aldridge, A., J. Krywy, E., S. Wirth, Ko A., Seybert, J., Winkler, coil,W . i,H .adJcb-anr C. Jacobs-Wagner, and C. H. Lim, B., W. Schofield, E. Emberly, and S. Saberi, E. Emberly, and S. Saberi, M. R. Losick, J. and , Z. D. and Rudner, B. Martinac, L., J. Hendel, R., A. Battle, M. J. T., Wood, Romantsov, and L. Stalker, C., Gill, E., D. Culham, S., Helbig, T., Romantsov, B. D. Weibel, and D. L. Renner, A. P. Boer, de and M. D. Raskin, enr .D n ebl .B. D. Weibel, and D. L. Renner, irsoy atra yaisa aestudy. case a as division. dynamics cell bacterial to microscopy: relation its and coli E. in clusters coli. Escherichia in Caulobacter. in division cell with segregation chromosome hogpt upxlpeiinaayi fbceilmrhgnssand morphogenesis bacterial of analysis precision subpixel throughput, coli. Escherichia in system .A,Oiir,V,Egl . ikno,A .adBara and J. A. Wilkinson, A., Engel, V., Olivieri, A., S. chemotaxis. dynamics. spatio-temporal intracellular n h oiinn faplrlclzto factor. localization polar a of integrity positioning membrane outer the for and essential is complex Tol-Pal caulobacter The (2010). eidct fpoencutr ngoigbacteria. growing in clusters protein of periodicity regulatory and structural bacterial of positioning polar proteins. the for factor localization fluorescent surface. yellow N-Terminal the bacterial of tag. artifact protein entire an is the MC1000/pLE7 coli Escherichia over occurs IcsA old bacterial of the to Microbiol. cleavage directly by targeted IcsP is determined IcsA pole: protein DivIVA surface Shigella protein unipolar The division cell subtilis microscopy. Bacillus electron transmission the of structure process. Z. proteins. localize fetplrcrmsm ergto nsouaigBclu subtilis. Bacillus sporulating in segregation chromosome Microbiol. polar effect mycobacteria. M. in Microbiol. apparatus secretory Mol. K. ESX-1 virulence-associated and Derbyshire, the of and coli proteins A. Escherichia T. Gray, in aggregation ageing. protein cellular and control quality protein B. of on consequences Bukau, and features A. Mogk, spatio-temporal growth S., A. apical Frangakis, M., of oligomerization Weiss, and function vivo in Streptomyces coelicolor. the in DivIVA determinant in involved Domains dynamics. division and patterns cells oscillation coli Min Escherichia affect Y-shaped in lengths branch mechanism: detection geometry oriainadrglto fbceilcrmsm ergto yaisby dynamics segregation proteins. chromosome localized bacterial polarly of regulation and coordination cells. bacterial in bacteria. in proteins the Biol. Perspect. of Harb. MscL. Spring comparison and Cold MscS, cells: AqpZ, coli ProW, LacY, Escherichia Bacteriol. ProP, J. proteins in membrane localization cytoplasmic Protein (2010). ProP transporter coli. osmosensory Escherichia of in localization polar promotes Cardiolipin (2007). coli. Escherichia in formation plane Chem. division regulate and phospholipids nhr h hoooesgeainadceoatcmcieyt the to machinery chemotactic and K. segregation M. Waldor, pole. chromosome and bacterial M. the B. Davis, anchors J., G. Jensen, A., eaieycre ein fEceihacl membranes. coli USA Escherichia of regions curved negatively coli. Escherichia of middle USA the Sci. to division directing for required 21) albce hoooesgeaini nodrdmultistep ordered an is segregation chromosome Caulobacter (2010). 108 287 6264-6269. , rc al cd c.USA Sci. Acad. Natl. Proc. rc al cd c.USA Sci. Acad. Natl. Proc. 96 32 49 38835-38844. , .Bacteriol. J. ora fCl cec 21)17 11 doi:10.1242/jcs.138628 11–19 127, (2014) Science Cell of Journal 4971-4976. , 192 MOJ. EMBO 367-377. , 1463-1475. , ee Dev. Genes 83 912-924. , Science 654-664. , LSONE PLoS o.Microbiol. Mol. o.Microbiol. Mol. LSCmu.Biol. Comput. PLOS ,E,Muchova E., ´, 29 20) tcatcasml fceoeetrclusters chemoreceptor of assembly Stochastic (2008). 21) pta oto ftecl iiinst yteMin the by site division cell the of control Spatial (2013). 194 i,L,Pugalr . aemn,U,Sujk V., Sourjik, U., Haselmann, S., Pruggnaller, L., nig, ¨ 2724-2733. , MOJ. EMBO 326 21) o-qiiru oa oaiaino proteins of localization polar Non-equilibrium (2013). 20) aAadteSjSoJsse obn to combine system Soj-Spo0J the and RacA (2003). nio.Microbiol Environ. 26 21) hoooedie pta atrigof patterning spatial driven Chromosome (2010). 6382-6386. , 21) rti uclua oaiaini bacteria. in localization subcellular Protein (2010). 8 1225-1228. , 2348-2360. , e64075. , 19) ai oet-oeoclaino protein a of oscillation pole-to-pole Rapid (1999). 20) iZ pta euao coordinating regulator spatial a MipZ, (2006). 21) iDadMn neatwt anionic with interact MinE and MinD (2012). 21) adoii irdmislclz to localize microdomains Cardiolipin (2011). 2 o.Microbiol. Mol. 68 64 21) pta raiaini bacterial in organization Spatial (2010). a000307. , 21) h eia rBctseeo in MreB helical The (2012). 29 107 ,K,Geoii . utg . Mu A., Lustig, M., Gregorini, K., ´, 99 1228-1236. , 1455-1465. , 21) oa sebyadscaffolding and assembly Polar (2012). 3068-3081. , o.Microbiol. Mol. 13831-13836. , 20) h i ytma eea cell general a as system Min The (2008). 20) oiinn fchemosensory of Positioning (2007). 14194-14198. , 6 21) mg nlssi fluorescence in analysis Image (2012). e1000986. , ESMcoil Lett. Microbiol. FEMS l . aeo,D . Briegel, E., D. Cameron, A., ll, ¨ . Bioessays 20) h n o bacteria how and Why (2009). .Bacteriol. J. hs e.Lett. Rev. Phys. MOJ. EMBO 52 .Bacteriol. J. 20) dniiaino a of Identification (2002). 21) utdmi hub multidomain A (2012). 1281-1290. , 80 k I.´k, 21) uniaieand Quantitative (2010). 20) Self-organized (2008). 612-627. , MOJ. EMBO rc al cd Sci. Acad. Natl. Proc. 34 26 Cell 21) elcycle Cell (2010). 427-436. , 192 20) Oligomeric (2004). 1615-1623. , rc al Acad. Natl. Proc. 190 ¨rdh, K. 126 4847-4858. , 297 101 21) High- (2011). 2106-2117. , 29 147-162. , 910-923. , 101-109. , 218101. , (1999). .Biol. J. (2009). ller, ¨ Mol. Mol. 19

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