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-2.0998 TD [(16)15.5(H)12.4(B)0(E)12.4(1)0(4)21.2(o)0(-)-447(c)0(e)14.7(l)0(l)16.5(s)0(,)-440.5(C)0(a)16.1(l)0(u)13.2(-)0(3)-447(c)0(e)20.4(l)0(l)16.5(s)-457.2(an)20.8(d)-456(M)0(a)18.9(d)0(i)13.2(n)0(-)19.6(D)0(a)16.9(r)0(b)19.6(y)-456(ca)14.7(-)]TJ -1.0015 -1.0528 TD [(ni)18.9(ne)-514.1(ki)18.9(dn)21.2(ey)-514.1(\(M)17.7(DC)12.6(K)12.4(\))-520.2(ep)15.1(it)16.5(he)20.8(li)16.5(al)-510.8(ce)14.7(ll)16.5(s)-519.8(w)0(e)16.9(r)0(e)-510(c)0(u)15.1(l)0(t)16.5(u)0(r)19.6(e)0(d)-514.1(a)0(s)]TJ 0 -1.0471 TD [(pr)19.6(ev)15.1(io)18.9(us)14.4(l)13.9(y)-387.7(de)20.8(sc)13.9(ri)17.3(be)20.8(d)-387.7(i)0(n)-373.8(f)0(u)19.6(l)0(l)-376.2([)0(1)13.9(7].)-384.8(In)-395.8(the)-397.7(presen)-8.2(t)-390.1(studi)-9.8(es,)]TJ 0 -1.0527 TD [(MDCK)-529.7(cell)-8.6(s)-519.8(wer)-8.2(e)-524.7(used)-528.2(as)-521(an)-525.5(experi)-9.4(mental)-531.1(refere)-10.6(nce)]TJ 0 -1.0471 TD [(because)-383.7(of)-378.7(thei)-8.2(r)-377.9(highly)-384.9(characteri)-11.9(zed)-378.7(inte)-8.2(rcellular)-388.3(adhe-)]TJ 0 -1.0528 TD [(sions.)-270.7(16HBE)-10.9(14o-)-266.4(cell)-8.6(s)-269.4(have)-265.6(the)-272.5(charact)-10.9(eristics)-274.1(of)-270.6(super-)]TJ 0 -1.0471 TD [(ficial)-298.6(epitheli)-10.7(al)-288.8(cell)-8.6(s)-286.5(and)-292.9(wer)-8.2(e)-285.7(origi)-10.2(nally)-293.5(deriv)-8.2(ed)-286.5(b)-6.5(y)-285.3(viral)]TJ 0 -1.0528 TD [(transform)-13.1(ation)-265.1(of)-264.9(cells)-266.6(dissociat)-11.5(ed)-258(from)-271.2(human)-263.4(bronchi)-9.7(al)]TJ 0 -1.0471 TD [(epithe)-10.1(lium)-437.8([17].)-444.8(Previou)-8.9(s)-434.4(phenot)-7.4(yping)-438.7(of)-441.3(Calu-3)-440.5(cell)-8.6(s,)]TJ 0 -1.0527 TD [(which)-255.2(are)-249.4(deriv)-8.2(ed)-246.6(from)-254.1(an)-246.6(ade)-8.8(nocarcinom)-10.3(a,)-246.2(sugges)-7.4(ts)-249.8(that)]TJ 0 -1.0471 TD [(they)-335.9(have)-339.6(seromuc)-12.1(ous)-333.5(attribut)-11.9(es)-333.2([18].)]TJ 1.0015 -1.0528 TD [(In)-190.9(ex)15.1(pe)20.8(ri)17.3(me)12(nt)18.9(s)-201.1(t)0(o)-191.7(e)0(x)20.8(a)0(m)12(i)0(n)18.9(e)-200.3(th)18.9(e)-200.3(b)0(r)14(e)0(a)20.4(k)0(d)15.5(o)0(w)17.3(n)-199.9(an)15.1(d)-199.9(f)0(o)19.6(r)0(m)16.5(a)0(t)18.4(i)0(o)13.2(n)]TJ -1.0015 -1.0471 TD [(of)-276.3(TJ)15.9(s,)-275.5(ce)20.4(ll)16.5(s)-286.5(w)0(e)16.9(r)0(e)-276.7(s)0(e)13.9(e)0(d)20.8(e)0(d)-275.1(a)0(t)-277.5(c)0(o)15.1(n)0(f)19.6(l)0(u)18.9(e)0(n)15.1(t)-287.7(de)20.8(ns)20(it)16.5(y)-285.3(i)0(n)18.8(t)0(o)-277.1(6)0(0)15.5(-)0(m)16.5(m)]TJ 0 -1.0527 TD [(P)11.4(e)0(t)12.8(r)0(i)-187.6(d)0(i)18.9(s)0(h)20(e)0(s)-190.9(c)0(o)20.8(n)0(t)18.9(a)0(i)12.8(n)0(i)18.9(n)0(g)-183.6(g)0(l)13.2(a)0(s)19.6(s)-195.4(co)15.1(ve)20.8(rs)18.4(li)16.5(ps)14.4(.)-193.9(C)0(e)16.1(l)0(l)16.5(s)-195.4(we)16.9(re)-185.6(al)18.4(lo)13.2(w)12.4(e)0(d)]TJ 0 -1.0471 TD [(to)-470.5(ad)15.1(he)15.1(re)-470.2(in)-470.5(no)15.5(rm)16.5(al)-470.9(cu)15.1(lt)16.5(ur)19.6(e)-484.9(m)0(e)17.7(d)0(i)18.9(u)0(m)-477(f)0(o)19.6(r)-480.4(5)-484.5(h)-478.8(an)15.1(d)-478.8(t)0(h)18.8(e)0(n)]TJ 0 -1.0527 TD [(wa)16.9(sh)20(ed)-371.8(an)20.8(d)-376.3(p)0(l)18.9(a)0(c)14.7(e)0(d)-366.1(i)0(n)-368.1(c)0(u)20.8(l)0(t)16.5(u)0(r)19.6(e)-376.7(me)12(di)18.9(um)-368.9(co)20.8(nt)18.9(ai)12.8(ni)18.9(ng)-365.7(5)-376.3(m)0(M)]TJ T* [(et)18.4(hy)15.5(le)18.4(ne)-240.9(gl)18.9(yc)15.1(ol)-237.2(te)12.8(tr)17.3(a-)19.2(ac)14.7(et)18.4(ic)-237.6(ac)14.7(id)-242.9(\(E)21.2(GT)87.2(A\))-240.4(to)-242.9("d)18.3(ow)17.3(n)-251.1(s)0(w)16.1(i)0(t)16.5(c)0(h)20.8(")]TJ 0 -1.0471 TD [(ex)15.1(tr)17.3(ac)20.4(el)18.5(lu)13.2(l)13.9(a)0(r)-140.1(C)11.6(a)]TJ 7.472 0 0 6.641 118.431 406.431 Tm (2+)Tj 9.963 0 0 9.963 126.142 402.009 Tm [(.)-142.6(T)0(h)17.1(r)0(e)13.5(e)-143.4(da)20.8(ys)-133.6(af)13.5(te)18.4(r)-144.6(p)0(l)18.9(a)0(t)18.4(i)0(n)18.9(g)0(,)-137.7(c)0(e)20.4(l)0(l)16.5(s)-144.2(we)16.9(re)-134.4(wa)16.9(sh)14.4(-)]TJ -6.8116 -1.0528 TD [(ed)-491.3(in)-493.3(fr)18(es)19.6(h)-501.5(m)0(e)17.7(d)0(i)18.9(u)0(m)-494.1(a)0(n)15.1(d)-495.8(ex)15.1(tr)17.3(ac)14.7(el)18.4(lu)18.9(la)18.4(r)-503.1(C)11.6(a)]TJ 7.472 0 0 6.641 229.039 396 Tm (2+)Tj 9.963 0 0 9.963 241.852 391.521 Tm [(re)19.2(st)17.7(or)13.9(ed)-491.3(to)]TJ -18.426 -1.0471 TD [(no)15.5(rm)16.5(al)-249(le)18.4(ve)15.1(ls)-249.8(\(")22.4(up)-251.9(sw)16.1(it)16.5(ch)20.8("\))16.7(.)-256.5(A)0(t)-252.5(v)0(a)15.1(r)0(i)17.3(o)0(u)21.2(s)-258(ti)16.5(me)17.7(s)-263.7(a)0(f)19.2(t)0(e)18.4(r)-258.4(th)18.9(e)-262.9(")13.4(up)]TJ 0 -1.0527 TD [(sw)16.1(it)16.5(ch)20.8(")-447.6(o)0(f)-435.6(C)0(a)]TJ 7.472 0 0 6.641 115.313 375.08 Tm (2+)Tj 9.963 0 0 9.963 123.024 370.601 Tm [(,)-444.2(c)0(e)14.7(l)0(l)16.5(s)-445.8(we)16.9(re)-436(fi)17.3(xe)20.8(d)-444.6(a)0(n)15.1(d)-444.6(th)18.9(ei)18.4(r)-446.2(T)0(J)-439.3(p)0(r)19.6(o)0(t)18.9(e)0(i)18.4(n)0(s)]TJ -6.4986 -1.0471 TD [(ex)15.1(am)17.7(in)18.9(ed)-314.9(by)-314.5(fl)17.3(uo)15.5(re)19.2(sc)13.9(en)20.8(t)-327.5(a)0(n)20.8(t)0(i)16.5(b)0(o)21.2(d)0(y)-320.2(l)13.9(ab)15.1(el)18.4(li)16.5(ng)15.5(.)]TJ /F4 1 Tf 0 -2.703 TD [(Measu)-7.8(r)32.2(emen)-7.8(t)-333.2(o)0(f)-334(perm)-8.6(eabilit)-9.6(y)]TJ /F3 1 Tf 1.0015 -1.8494 TD [(Me)13.2(as)19.6(ur)19.6(em)12(en)20.8(ts)-335.2(of)-333.2([)]TJ 7.472 0 0 6.641 142.809 319.238 Tm (14)Tj 9.963 0 0 9.963 150.123 314.816 Tm [(C)11.6(])0(-)12.4(m)0(a)17.7(n)0(n)21.2(i)0(t)16.5(o)0(l)-334(c)0(l)12.8(e)0(a)20.4(r)0(a)19.2(n)0(c)15.1(e)-342.6(an)15.1(d)-342.2(p)0(e)20.8(r)0(m)10.8(e)0(-)]TJ -9.2187 -1.0528 TD [(ab)15.1(il)22.2(it)16.5(y)-495.8(w)0(e)16.9(r)0(e)-487.2(p)0(e)15.1(r)0(f)18(o)0(r)19.6(m)0(e)17.7(d)-495.8(ex)15.1(ac)20.4(tl)16.5(y)-495.8(a)0(s)-486.8(d)0(e)20.8(s)0(c)13.9(r)0(i)17.3(b)0(e)20.8(d)-495.8(pr)13.9(ev)20.8(io)18.9(us)14.4(ly)]TJ 0 -1.0471 TD [([1)19.6(9].)-458.7(Exp)-6.4(eriments)-472.8(wer)-8.2(e)-467.8(conducte)-9.7(d)-461.7(i)0(n)-470.5(confluen)-10.9(t)-464.1(mono-)]TJ 0 -1.0528 TD [(layer)-8.6(s)-371.8(o)0(f)-373(epitheli)-10.7(al)-374.2(cells)-374.8(whi)-8.6(ch)-371.8(had)-372.6(been)-379.4(propaga)-7.7(ted)-375(on)]TJ 0 -1.0471 TD [(ultrathi)-11.9(n)-228.4(underco)-7.7(ats)-228.2(of)-230.7(ungell)-9.7(ed)-229.6(Matrigel)-239.5(in)-225.8(Cos)-8.2(tar)-228.6(T)29.3(rans-)]TJ 0 -1.0528 TD [(wells)-475(\(Corni)-9.2(ng-Cost)-9.5(ar)-470.2(Ltd.,)-476.2(High)-470.3(W)49.3(ycomb)-7.4(e,)-473.9(Bucking-)]TJ 0 -1.0471 TD [(hams)-7.8(hire,)-309.4(UK\).)-308(All)-306.9(cell)-310.2(lines)-306.1(wer)-8.2(e)-302.8(seeded)-314.3(at)-305.9(densities)-313.1(of)]TJ 0 -1.0527 TD 0 Tw (5)Tj /F9 1 Tf 0.5008 0 TD (3)Tj /F3 1 Tf 0.8365 0 TD (10)Tj 7.472 0 0 6.641 81.581 245.99 Tm (5)Tj 9.963 0 0 9.963 88.611 241.568 Tm [(cell)-8.6(s)]TJ /F10 1 Tf 1.8096 0.1992 TD (.)Tj /F3 1 Tf 0.2675 -0.1992 TD [(inser)-9.4(t)]TJ 7.472 0 0 6.641 131.131 245.99 Tm (-1)Tj 9.963 0 0 9.963 137.31 241.568 Tm (.)Tj /F4 1 Tf -7.9326 -2.703 TD [(Measu)-7.8(r)32.2(emen)-7.8(t)-333.2(o)0(f)-334(trans)-8.6(epitheli)-10.7(al)-334(electrica)-11.1(l)-333.2(r)26.5(esistance)]TJ /F3 1 Tf 1.0015 -1.8494 TD [(T)40.6(r)0(a)13.5(n)0(s)20(e)0(p)15.1(i)13.9(th)13.2(et)18.4(hi)18.9(al)-214.9(el)18.4(ec)14.7(tr)17.3(ic)18.4(al)-214.9(re)13.5(si)17.7(st)17.7(an)20.8(ce)-218.6(\(T)21.2(ER)12.4(\))-224.3(w)12.4(as)-219.4(me)17.7(as)19.6(ur)25.3(-)]TJ -1.0015 -1.0527 TD [(ed)-286.5(in)-282.7(En)17.1(do)15.5(hm)]TJ 7.472 0 0 6.641 114.066 190.205 Tm 0.0001 Tc (TM)Tj 9.963 0 0 9.963 128.013 185.726 Tm 0 Tc [(ti)16.5(ss)13.2(ue)-280.8(re)13.5(si)17.7(st)17.7(an)20.8(ce)-286.9(me)12(as)19.6(ur)19.6(em)12(en)20.8(t)-293.4(c)0(h)15.1(a)0(m)17.7(b)0(e)15.1(r)0(s)]TJ -6.9994 -1.0471 TD [(\(W)86.8(or)19.6(ld)-237.2(Pr)14.7(ec)20.4(is)12(i)13.9(o)0(n)-240.5(I)0(n)13.9(s)0(t)17.7(r)0(u)19.6(m)0(e)17.7(n)0(t)18.9(s)0(,)-241.3(I)0(n)19.6(c)0(.)15.5(,)-245.1(Sa)15.9(ra)19.2(so)14.3(ta)18.4(,)-250.8(F)11.4(L,)-238.6(US)18.1(A\))-240.4(by)]TJ 0 -1.0528 TD [(ap)15.1(pl)18.9(yi)18.9(ng)-269(a)-280(\213)11.8(20)15.5(-)]TJ /F11 1 Tf 6.2368 0 TD (m)Tj /F3 1 Tf 0.5634 0 TD [(A)-277.8(s)0(q)14.4(u)0(a)20.8(r)0(e)-271(w)0(a)16.9(v)0(e)-269.4(a)0(l)18.5(t)0(e)12.8(r)0(n)19.6(a)0(t)18.4(i)0(n)18.9(g)-279.6(cu)15.1(rr)18(en)20.8(t)-282(a)0(c)20.4(r)0(o)13.9(s)0(s)]TJ -6.8002 -1.0471 TD [(th)18.9(e)-234.5(m)0(o)18.1(n)0(o)21.2(l)0(a)12.8(y)0(e)20.8(r)0(s)-226.2(a)0(t)-220.6(1)0(2)21.2(.)0(5)-223.1(H)0(z)16.9(.)-233.7(T)12.2(h)0(e)-223.9(v)0(a)20.8(l)0(u)13.2(e)0(s)-219.4(o)0(f)-225.1(T)12.2(ER)-226.6(ci)18.5(te)18.4(d)-228.4(i)0(n)-225.8(t)0(h)18.9(i)0(s)]TJ 0 -1.0528 TD [(ma)17.7(nu)15.5(sc)19.6(ri)17.3(pt)-197.4(we)16.9(re)-197(ma)12(de)-195.4(wi)14.9(th)-197.4(5)-205.6(m)13.2(L)-209.7(m)0(e)17.7(d)0(i)18.9(u)0(m)-198.2(i)0(n)-197.4(t)0(h)18.9(e)-206(ou)15.5(te)18.4(r)-207.2(w)0(e)16.9(l)0(l)]TJ 0 -1.0471 TD [(an)15.1(d)-228.4(0)0(.)21.6(1)-234.1(mL)-225(me)17.7(di)18.9(um)-220.9(in)-225.8(th)18.9(e)-228.8(T)40.6(ra)13.5(ns)20(we)16.9(ll)-222.5(in)18.9(se)13.9(rt)17.3(.)-228(T)0(h)17.1(e)-228.8(re)13.5(si)17.7(st)17.7(an)20.8(ce)]TJ 0 -1.0528 TD [(of)-464.1(ce)14.7(ll)16.5(-f)18(re)19.2(e)-473.5(T)40.6(ra)13.5(ns)20(we)16.9(ll)-467.2(in)18.9(se)19.6(rt)17.3(s)-474.3(w)0(a)16.9(s)-474.3(me)17.7(as)13.9(ur)19.6(ed)-468.6(fo)19.6(r)-474.7(c)0(o)20.8(n)0(t)13.2(r)0(o)19.6(l)]TJ 0 -1.0471 TD [(pu)15.5(rp)19.6(os)20(es)-316.1(an)15.1(d)-325.1(p)0(r)19.6(o)0(v)15.5(e)0(d)-314.9(t)0(o)-311.2(b)0(e)-314.9(m)0(i)15.7(n)0(i)18.9(m)0(a)12(l)-321.8(\()]TJ /F11 1 Tf 14.784 0 TD (<)Tj /F3 1 Tf 0.5463 0 TD (1)Tj /F12 1 Tf 0.8251 0 TD (V)Tj /F10 1 Tf 0.7056 0.1992 TD (.)Tj /F3 1 Tf 0.2731 -0.1992 TD (cm)Tj 7.472 0 0 6.641 240.831 116.957 Tm (2)Tj 9.963 0 0 9.963 244.46 112.535 Tm (\).)Tj /F4 1 Tf 6.51 66.1583 TD [(Cel)-9.8(lular)-351.2(staining)-355.9(by)-349.1(fluor)30.1(escent)-356.5(pr)31.4(obe)-8.4(s)-343.4(and)-349.4(indi)-7.8(r)32.2(ect)-352.6(im-)]TJ 0 -1.0641 TD [(muno)-7(cytoch)-8.2(emistr)-10.4(y)]TJ /F3 1 Tf 1.0015 -2.4469 TD [(T)69.1(o)-256.8(de)20.8(te)12.8(rm)16.5(in)18.8(e)-257.2(t)0(h)18.9(e)-257.2(ex)20.8(te)18.4(nt)-248.6(of)-247.8(ce)20.4(ll)-251(co)20.8(ve)15.1(ra)19.2(ge)-246.6(ov)15.5(er)-248.2(T)40.6(r)0(a)19.2(n)0(s)20(w)0(e)16.9(l)0(l)]TJ -1.0015 -1.0641 TD [(in)13.2(se)19.6(rt)17.3(s)-349.1(a)0(n)15.1(d)-347.9(th)18.8(e)-348.3(v)0(i)18.9(a)0(b)15.1(i)0(l)16.5(i)13.9(ty)-339.7(of)-344.6(ce)20.4(ll)16.5(s)-349.1(a)0(t)-340.1(a)0(n)15.1(y)-347.9(ti)16.5(m)13.2(e)-354(po)21.2(in)18.9(t)-350.3(d)0(u)15.5(r)0(i)17.3(n)0(g)]TJ 0 -1.0584 TD [(cu)15.1(lt)16.5(ur)19.6(e,)-212.1(sa)19.6(mp)18.1(le)-214.9(i)13.9(n)0(s)14.3(e)0(r)19.2(t)0(s)-210(w)0(e)16.9(r)0(e)-214.1(s)0(e)19.6(l)0(e)18.4(c)0(t)18.5(e)0(d)-212.5(a)0(t)-209.2(r)0(a)13.5(n)0(d)21.2(o)0(m)-209.5(a)0(n)15.1(d)-217(st)17.7(ai)18.5(ne)15.1(d)]TJ 0 -1.0641 TD [(wi)14.9(th)-345.3(et)18.4(hi)13.2(di)18.9(um)-346.1(br)19.6(om)18.1(id)13.2(e)-354(a)0(n)20.8(d)-353.6(ac)14.7(ri)17.3(di)18.9(ne)-349.1(or)19.6(an)15.1(ge)-343.4(\(M)17.7(er)13.5(ck)-343.4(Lt)20.4(d.)15.9(,)]TJ T* [(Lu)17.1(tt)16.5(er)13.5(w)12.4(o)0(r)13.9(t)0(h)18.8(,)-461.3(L)12.2(e)0(i)18.5(c)0(e)14.7(s)0(t)17.7(e)0(r)19.2(s)0(h)14.4(i)0(r)17.3(e)0(,)-451.1(U)12.4(K\))15.7(.)-461.3(T)0(h)17.1(e)-462.1(ce)20.4(ll)16.5(s)-457.2(w)0(e)11.2(r)0(e)-447.4(r)0(i)17.3(n)0(s)14.3(e)0(d)]TJ 0 -1.0584 TD [(tw)14.9(ic)18.4(e)-240.2(w)0(i)14.9(t)0(h)-231.5(p)0(h)21.2(o)0(s)14.4(p)0(h)21.2(a)0(t)12.8(e)0(-)19.2(b)0(u)15.5(f)20.4(fe)19.2(re)13.5(d)-239.8(s)0(a)19.6(l)0(i)16.5(n)0(e)-229.6(\()0(P)14.7(B)11.6(S\))-230(an)15.1(d)-239.8(1)0(0)15.5(0)]TJ /F11 1 Tf 20.5827 0 TD (m)Tj /F3 1 Tf 0.5634 0 TD 0.2382 Tc [(Lo)238.2(f)]TJ -21.1461 -1.0641 TD 0 Tc [(ac)14.7(ri)17.3(di)18.9(ne)-235.2(or)14(an)20.8(ge)-235.2(an)15.1(d)-245.5(1)0(0)21.2(0)]TJ /F11 1 Tf 9.6455 0 TD (m)Tj /F3 1 Tf 0.5634 0 TD -0.0189 Tc [(L)-268.5(e)-18.9(th)-18.9(id)-18.9(iu)-18.9(m)-256.9(b)-18.9(ro)-18.9(m)-6.5(i)-18.9(de)-264.8(\()-18.9(1)-255.3(m)-18.9(g)]TJ /F10 1 Tf 10.3568 0.1992 TD 0 Tc (.)Tj /F3 1 Tf 0.2731 -0.1992 TD (mL)Tj 7.472 0 0 6.641 530.419 677.594 Tm (-1)Tj 9.963 0 0 9.963 309.316 662.57 Tm [(fo)13.9(r)-195.8(e)0(a)14.7(c)0(h)15.1(\))-195.8(ad)15.1(de)20.8(d)-199.9(u)0(n)15.5(d)0(e)20.8(r)-201.5(su)20(bd)15.5(ue)20.8(d)-199.9(i)0(l)16.5(l)0(u)18.9(m)0(i)15.7(n)0(a)15.1(t)0(i)16.5(o)0(n)21.2(.)-199.6(Th)17.1(e)-200.3(c)0(e)20.4(l)0(l)16.5(s)-201.1(we)16.9(re)]TJ 0 -1.0584 TD [(im)15.7(me)12(di)18.9(at)18.5(el)18.5(y)-325.1(e)0(x)15.1(a)0(m)17.7(i)0(n)18.8(e)0(d)-314.9(b)0(y)-314.5(f)0(l)17.3(u)0(o)15.5(r)0(e)19.2(s)0(c)19.6(e)0(n)15.1(c)0(e)-315.3(m)0(i)15.7(c)0(r)19.2(o)0(s)14.4(c)0(o)20.8(p)0(y)-314.5(u)0(s)14.4(i)0(n)18.8(g)]TJ 0 -1.0641 TD [(a)-285.7(Z)0(e)16.7(i)0(s)17.7(s)-286.5(Ax)17.3(io)18.8(ve)15.1(rt)-273(10)-274.7(mi)15.7(cr)19.2(os)14.4(co)20.8(pe)-275.1(\(C)14.9(ar)19.2(l)-287.7(Z)0(e)16.7(i)0(s)17.7(s)-280.8(Lt)14.7(d.)15.9(,)-279.2(W)77.7(el)18.4(wy)17.3(n)]TJ T* [(Ga)16.9(rd)13.9(en)-235.2(ci)18.5(ty)75.8(,)-245.1(H)12.4(er)13.5(tf)17.3(or)19.6(ds)14.4(hi)18.9(re)19.2(,)-245.1(U)0(K)13.4(\))-247.1(w)12.4(i)0(t)16.5(h)-245.5(bl)18.9(ue)15.1(/g)18.9(re)13.5(en)-235.2(ex)20.8(ci)12.8(ta)18.4(ti)16.5(on)]TJ 0 -1.0584 TD [(fi)17.3(lt)16.5(er)-316.5(se)13.9(t)13.9(s)0(.)]TJ 1.0015 -1.0527 TD [(In)13.9(te)18.4(rc)19.2(el)12.8(lu)18.9(la)18.4(r)-514.5(j)0(u)13.2(n)0(c)20.8(t)0(i)16.5(o)0(n)15.5(s)-514.1(we)16.9(re)-510(ex)20.8(am)12(i)13.9(n)0(e)15.1(d)-518.6(by)-502.3(fl)17.3(uo)15.5(re)19.2(sc)13.9(en)20.8(t)]TJ -1.0015 -1.0641 TD [(an)15.1(ti)16.5(bo)21.2(dy)-456.8(la)18.4(be)20.8(ll)16.5(in)13.2(g)-461.7(u)0(s)20(i)0(n)18.8(g)-461.7(mo)18.1(no)15.5(cl)18.4(on)15.5(al)-453.9(an)20.8(ti)16.5(bo)15.5(di)18.9(es)-452.7(\(m)16.5(Ab)17.3(s\))]TJ 0 -1.0584 TD [(fo)13.9(r)-440.5(o)0(c)15.1(c)0(l)18.5(u)0(d)15.5(i)13.9(n,)-433.6(E-)15.5(ca)14.7(dh)21.2(er)13.5(i)13.9(n)0(,)-433.6(a)0(n)15.1(d)-444.6(de)20.8(sm)16.9(op)15.5(la)18.4(ki)18.9(n)-444.6(a)0(s)-429.9(d)0(e)15.1(s)0(c)19.6(r)0(i)17.3(b)0(e)15.1(d)]TJ 0 -1.0641 TD [(el)12.8(se)19.6(wh)17.3(er)19.2(e)-484.9([)0(1)13.9(6].)-487.2(The)-496.2(mAbs)-500.7(used)-499.7(were)-498.8(rat)-496.1(antio)-9(ccludin)]TJ T* [(\(MO)-9.8(C37;)-465.4(a)-462.1(kind)-466.4(gift)-464.7(of)-464.1(S.)-461.3(T)57.7(sukita,)-466.6(Kyoto)-471.1(Univers)-9.1(ity)61.2(,)]TJ 0 -1.0584 TD [(Kyot)-9.4(o,)-285.7(Japan\))-292.7([20],)-291.2(rat)-291.2(anti)-8.2(-E-cadher)-11.3(in)-288.4(\(DECM)-8.8(A-1;)-289.8(Sig-)]TJ 0 -1.0641 TD [(ma,)-278.6(Po)-6.4(ole,)-272.9(Dor)-7.8(set,)-279.8(UK\))-279.2([21])-273.7(an)-7.6(d)-273.9(mouse)-276(an)-7.6(tidesmo)-9.1(plakin)]TJ T* [(\(1)25.3(1-5F\))-353.3([22].)-353.7(In)-350.3(human)-354.5(cell)-8.6(s)-349.1(ZO-1)-354.1(was)-354.9(labelled)-355.9(using)-354.5(a)]TJ 0 -1.0584 TD [(rabbi)-8.2(t)-333.2(polyclonal)-339.9(anti)-8.2(body)-333.1(\(a)-333.6(kind)-335.5(gift)-333.8(from)-339.4(T)74.8(.)-336.1(Fleming,)]TJ 0 -1.0641 TD [(Uni)-8.6(versity)-411.6(of)-412.8(Southampto)-9.5(n,)-405.2(So)-6.4(uthampto)-8.7(n,)-405.2(UK)-9.4(\),)-406.8(and)-406.7(in)]TJ T* [(MD)-7.4(CK)-357.3(cells)-363.4(with)-358.9(rat)-359.5(mAb)-356.5(R40)-7(.76)-354.7(\(a)-356.3(kind)-358.2(gift)-362.2(from)-362.2(B.)]TJ 0 -1.0584 TD [(Stevens)-8.9(on,)-525.4(Uni)-8.6(versity)-531.1(of)-521(Al)-7.8(berta,)-526.8(Edmont)-8.5(on,)-525.4(Albert)-9.6(a,)]TJ 0 -1.0641 TD [(Can)-7.4(ada\))-250.2([23].)]TJ /F11 1 Tf 5.6962 0 TD (b)Tj /F3 1 Tf 0.552 0 TD [(-C)-7.8(atenin)-249.2(was)-246.8(label)-9.4(led)-244.1(using)-246.4(a)-245.9(rabbi)-8.2(t)-242.2(poly-)]TJ -6.2482 -1.0641 TD [(clonal)-289(antibody)-284.4(\(Sigm)-10.2(a\).)-282.7(Antibod)-8.3(ies)-279.4(wer)-8.2(e)-274.3(dilu)-7.8(ted)-283.9(for)-278.6(use)]TJ 0 -1.0584 TD [(in)-294.1(PBS/)-8.6(0.2%)-289.6(volum)-9.3(e\(v\)/v)-295.1(T)57.7(ween-20)-296.2(contai)-10.2(ning)-290(10%)-294.9(v/v)]TJ 0 -1.0641 TD [(non)-8(immune)-292.5(serum)-10.2(.)-284.9(A)0(l)-7.8(l)-287.7(of)-287.7(the)-295.3(cells)-289.4(used)-294.9(in)-288.4(the)-289.6(studi)-9.8(es)-287.6(of)]TJ T* [(inte)-8.2(rcellular)-479.3(junctions)-470.8(wer)-8.2(e)-467.8(fixed)-469.1(in)-470.5(ice-cold)-474.6(methanol)]TJ 0 -1.0584 TD [(and)-310(blocke)-7.8(d)-302.4(i)0(n)-311.2(10%)-306.3(v/v)-312(nonimmu)-8.3(ne)-303.5(serum)-317.4(before)-310.2(prim-)]TJ 0 -1.0641 TD [(ary)-300.2(antibody)-301.5(binding.)-300.7(For)-299(ZO-1)-297.2(and)-298.6(E-cadherin)-303.1(staining)-9.1(,)]TJ T* [(modi)-8.6(fied)-457(CSK)-454(buffer)-442.2(\(com)-8.2(position)-8.7(:)-452.7(NaCl)-458.3(50)-451.1(mM,)-450.9(su-)]TJ 0 -1.0584 TD [(crose)-309(3,000)-305(mM,)-308.6(piper)-8.2(azine-)]TJ /F4 1 Tf 11.603 0 TD (N)Tj /F3 1 Tf 0.6715 0 TD (,)Tj /F4 1 Tf 0.2504 0 TD (N)Tj /F8 1 Tf 0.6658 0 TD (')Tj /F3 1 Tf 0.2219 0 TD [(-bis)-304.1([2-eth)-10.6(anesulphoni)-9.5(c)]TJ -13.4126 -1.0641 TD [(acid])-355.7(\(PIPE)-9.6(S;)-344.6(pH)-352.6(6.8\))-351.4(10)-348.7(mM,)-354.1(MgCl)]TJ 7.472 0 0 6.641 461.537 385.909 Tm (2)Tj 9.963 0 0 9.963 468.737 387.553 Tm [(3)-347.9(mM,)-348.4(0.5%)-352.2(v/v)]TJ -16.0018 -1.0641 TD [(T)29.3(rit)-8.6(on)-348.7(X-1)-7.8(00,)-349(4-\(2-)-8.6(aminoethy)-9.9(l\)-ben)-10.6(zenesulphonyl)-364.2(fluor-)]TJ 0 -1.0584 TD [(ide)-249.8(hydroch)-7.3(loride)-250.3(1.2)-240.9(mM\))-248(was)-246.8(appli)-9(ed)-240.9(before)-253.3(fixation)-248.6(to)]TJ 0 -1.0641 TD [(remo)-8.2(ve)-223.9(staining)-225.4(not)-220.9(associ)-10.1(ated)-222.5(with)-222.3(cytos)-7.8(keletal)-224.2(compo)-7.4(n-)]TJ T* [(ents.)-508.2(Indir)-9.4(ect)-506.2(immunoc)-8.7(ytoche)-7.8(mistry)-510.3(was)-508.6(performed)-510.7(in)]TJ 0 -1.0584 TD [(speci)-8.2(mens)-406.1(label)-9.4(led)-403.4(with)-404.4(one)-401(or)-407.2(two)-401.2(prim)-10.2(ary)-402.6(antibodies)]TJ 0 -1.0641 TD [(dep)-8.4(ending)-326(on)-320.2(the)-323.7(ex)-7.6(periment)-10.7(.)-319.1(For)-327.5(single-antibo)-11.6(dy)-325.9(label-)]TJ T* [(ling)-406.2(studies)-411.2(the)-403.4(affinity-purified)-409.2(seconda)-7.7(ry)-401.5(immunog)-8.3(lo-)]TJ 0 -1.0584 TD [(buli)-7.8(n)-370.7(G)-380.3(was)-377.7(conjuga)-8.5(ted)-375(with)-375.9(fluor)-9.4(escei)-8.6(n)-370.7(isothioc)-9.1(yanate)]TJ 0 -1.0641 TD [(\(FIT)-9.6(C\).)-292.7(In)-299(dual)-296.1(label)-9.4(ling)-298(studies,)-297.8(an)-297.8(appropr)-8.9(iately)-297.1(paire)-8.6(d)]TJ T* [(combi)-9.7(nation)-191.8(of)-185.2(affinity-purified,)-199.1(immunoa)-8.7(bsorbe)-7.8(d)-182.9(second)]TJ 0 -1.0584 TD [(anti)-8.2(bodies)-492.2(conjugated)-491.6(wi)-7.8(th)-481.9(fluor)-9.4(escein)-493.1(dichlorot)-10.3(riazine)]TJ 0 -1.0641 TD [(\(DT)67.7(AF\))-246(and)-247.4(tetraethy)-11.1(lrhod)-7.8(amine)-249.2(isothioc)-9.1(yanate)-247.2(\(TRIT)-8.6(C\))]TJ T* [(was)-417.5(used)-414.4(\(Chemi)-11.1(con)-412.4(Internati)-12.3(onal,)-415.9(Harrow)52.3(,)-410.1(Middlesex)-9.9(,)]TJ 0 -1.0584 TD [(UK)-9.4(\).)-480.7(F-actin)-484(was)-485.8(label)-9.4(led)-483.1(using)-479.7(rhodam)-9.7(ine)-483.1(phall)-9(oidin)]TJ 0 -1.0641 TD [(\(C)-7.8(ambridge)-224.6(Bi)-8.6(oscience,)-227.9(Cambridge)-13.8(,)-216.6(UK\))-222.3(in)-220.2(cell)-8.6(s)-218.2(that)-224.5(had)]TJ T* [(bee)-8.8(n)-313.8(fixed)-321.2(in)-316.9(3.7%)-323.7(v/v)-317.7(formaldeh)-10.7(yde)-321.4(and)-315.7(perm)-8.2(eabilized)]TJ 0 -1.0584 TD [(in)-339.7(0.1%)-335.1(v/v)-334.7(T)29.3(rit)-8.6(on)-331.6(X-1)-7.8(00.)]TJ 1.0015 -1.0528 TD [(Ep)17.1(if)17.3(lu)13.2(or)19.6(es)19.6(ce)14.7(nc)20.8(e)-581.6(m)0(i)15.7(c)0(r)19.2(o)0(s)14.4(c)0(o)20.8(p)0(y)-570.6(w)0(a)16.9(s)-582.4(pe)20.8(rf)12.4(or)19.6(me)17.7(d)-581.2(u)0(s)20(i)0(n)18.8(g)-581.2(a)]TJ -1.0015 -1.0584 TD [(Ze)16.7(is)17.7(s)-360.5(A)12.4(xi)13.2(ov)21.2(er)19.2(t)-356(1)0(0)-348.7(m)0(i)15.7(c)0(r)19.2(o)0(s)14.4(c)0(o)20.8(p)0(e)-349.1(w)0(i)14.9(t)0(h)-345.3(o)0(i)18.9(l)-356(im)15.7(me)17.7(rs)12.8(i)13.9(o)0(n)-348.7(o)0(b)15.5(j)0(e)18.5(c)0(-)]TJ 0 -1.0641 TD [(ti)16.5(ve)15.1(s)-206.8(\()0(C)14.9(a)0(r)19.2(l)-208(Ze)16.7(is)17.7(s)-206.8(L)0(t)14.7(d)0(.)15.9(\))0(.)-196.2(F)11.4(lu)13.2(or)19.6(es)19.6(ce)14.7(nc)15.1(e)-206(e)0(x)15.1(c)0(i)18.5(t)0(a)18.4(t)0(i)16.5(o)0(n)-200.7(w)12.4(as)-202.3(at)-197.8(45)15.5(0\261)]TJ T* [(49)15.5(0)-279.6(n)0(m)-266.4(a)0(n)15.1(d)-273.9(em)12(i)13.9(s)0(s)13.2(i)0(o)18.9(n)-279.6(at)]TJ /F11 1 Tf 9.7081 0 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[(to)13.2(rt)17.3(uo)21.2(us)14.4(,)-347.5(t)0(h)18.8(e)-348.3(me)17.7(an)-343.4(ce)20.4(ll)-342(di)18.8(am)12(et)18.4(er)19.2(s)-349.1(s)0(u)14.4(g)0(g)21.2(e)0(s)13.9(t)-344.6(th)13.2(at)-340(th)18.9(e)-348.3(b)0(a)15.1(r)0(r)18(i)0(e)18.4(r)]TJ 0 -1.0528 TD [(pr)13.9(op)21.2(er)13.5(ti)16.5(es)-418.5(of)-424.2(mo)18.1(no)21.2(la)12.8(ye)20.8(rs)-425.4(fo)19.6(rm)16.5(ed)-423(by)-416.9(th)13.2(e)-428(t)0(h)18.8(r)0(e)13.5(e)-428(ce)14.7(ll)-421.7(li)16.5(ne)20.8(s)]TJ 0 -1.0471 TD [(sh)14.4(ou)21.2(ld)-316.9(be)-314.9(co)15.1(mp)18.1(ar)19.2(ab)15.1(le)18.4(.)]TJ 1.3771 -1.0528 TD [(F)11.4(i)0(g)18.9(u)0(r)14(e)0(s)-355.9(1)0(a)-349.1(and)-378.3(b)-370.7(show)-378.1(that,)-378.5(after)-380.1(plating)-377.1(on)-371.4(0)-6.5(.4-)]TJ /F11 1 Tf 20.0763 0 TD (m)Tj /F3 1 Tf 0.5747 0 TD (m)Tj -22.0281 -1.0471 TD [(porosi)-9.7(ty)-652.6(T)29.3(ranswe)-12.1(lls,)-651.7(monol)-9.3(ayers)-656.1(of)-657.5(16HBE14o)-7.5(-)-656.8(and)]TJ 0 -1.0528 TD [(Cal)-9.8(u-3)-254.3(cells)-255.3(develo)-7.8(ped)-253.1(incre)-8.6(asing)-258.2(TER)-254.3(values)-259.3(which)-255.2(rea-)]TJ 0 -1.0471 TD [(che)-8.8(d)-262.5(maxi)-9(ma)-266.8(of)-264.9(\(mea)-8.6(n\213)]TJ 6.974 0 0 6.974 406.035 522.879 Tm 0.0001 Tc (SEM)Tj 9.963 0 0 9.963 420.321 522.879 Tm 0 Tc [(\))-264.1(3)0(2)-7.2(7.8\21351.7)-271.6(and)-264.5(359)-8(.5\21346.7)]TJ /F12 1 Tf -11.1421 -1.0528 TD (V)Tj /F10 1 Tf 0.7227 0.1992 TD (.)Tj /F3 1 Tf 0.2731 -0.1992 TD (cm)Tj 7.472 0 0 6.641 331.257 516.812 Tm (2)Tj 9.963 0 0 9.963 338.57 512.39 Tm [(respective)-11.1(ly)-356.7(\(n)]TJ /F11 1 Tf 5.958 0 TD (=)Tj /F3 1 Tf 0.552 0 TD [(5\))-355.9(6\2618)-355.1(da)-7.6(ys)-355.5(after)-357.3(plat)-8.2(ing.)-357.9(There-)]TJ -9.4463 -1.0471 TD [(after)23.9(,)-313.4(TER)-316.8(declined)-319.4(to)-311.2(values)-316.2(of)-316.1(~120)]TJ /F12 1 Tf 15.5978 0 TD (V)Tj /F10 1 Tf 0.7227 0.1992 TD (.)Tj /F3 1 Tf 0.2675 -0.1992 TD (cm)Tj 7.472 0 0 6.641 486.652 506.381 Tm (2)Tj 9.963 0 0 9.963 493.512 501.959 Tm [(by)-308.8(day)-315.7(14.)]TJ -18.4886 -1.0527 TD [(In)-532.3(co)-7.6(ntrast,)-536.4(MDCK)-535.4(cell)-537.8(monola)-10.5(yer)-533.5(TER)-533.1(was)-531.3(5)-6.5(0\2137.5)]TJ /F12 1 Tf 0 -1.0471 TD (V)Tj /F10 1 Tf 0.7227 0.1992 TD (.)Tj /F3 1 Tf 0.2731 -0.1992 TD (cm)Tj 7.472 0 0 6.641 331.257 485.461 Tm (2)Tj 9.963 0 0 9.963 339.534 481.039 Tm [(on)-451.1(day)-452.3(1)-450.3(after)-459.7(plating)-456.7(and)-452.2(then)-455.4(decli)-9.4(ned)-452.2(to)-453.5(an)]TJ /F18 1 Tf 10 0 0 10 75.286 456.064 Tm 0 0 0 1 k /GS3 gs (a\))Tj /F19 1 Tf 6 0 0 6 277.181 360.91 Tm (l)Tj /F18 1 Tf 0 10 -10 0 95.786 370.511 Tm (TER )Tj /F20 1 Tf 1.868 0 TD (W)Tj /F18 1 Tf 0.768 0 TD (\341cm)Tj 0 8 -8 0 92.951 410.08 Tm (2)Tj ET 0 0 0 1 K 0 J 0 j 0.5 w 3.864 M []0 d /GS4 gs 1 i 120.368 451.683 m 120.368 334.711 l 120.632 451.491 m 117.89 451.491 l 120.632 427.759 m 117.89 427.759 l 120.632 404.665 m 117.89 404.665 l 120.632 381.743 m 117.89 381.743 l 120.632 357.756 m 117.89 357.756 l 120.632 334.79 m 117.89 334.79 l S BT 10 0 0 10 111.673 331.565 Tm /GS3 gs (0)Tj ET /GS4 gs 120.142 331.597 m 292.565 331.597 l 120.355 331.806 m 120.355 329.065 l 144.084 331.806 m 144.084 329.065 l 167.814 331.806 m 167.814 329.065 l 191.023 331.806 m 191.023 329.065 l 214.768 331.806 m 214.768 329.065 l 237.874 331.806 m 237.874 329.065 l 260.725 331.806 m 260.725 329.065 l 284.469 331.806 m 284.469 329.065 l S BT 10 0 0 10 102.833 354.431 Tm /GS3 gs (100)Tj 0 2.358 TD (200)Tj 0.0101 2.3087 TD (300)Tj 0 2.3248 TD (400)Tj 0 2.3493 TD (500)Tj /F19 1 Tf 6 0 0 6 270.237 363.9 Tm (l)Tj -5.7578 -0.2639 TD (l)Tj -1.9694 1.8751 TD (l)Tj -3.9032 4.7037 TD (l)Tj 1.963 -0.9815 TD (l)Tj -3.9414 2.1683 TD (l)Tj -5.8317 -11.1204 TD (l)Tj -1.9337 -1.6265 TD (l)Tj -1.9337 0.0293 TD (l)Tj ET /GS4 gs 283.065 366.425 m 286.055 366.425 l 283.065 360.184 m 286.055 360.184 l 284.648 366.249 m 284.648 360.184 l 271.019 373.193 m 274.009 373.193 l 271.019 358.777 m 274.009 358.777 l 272.602 373.193 m 272.602 358.777 l 236.472 370.731 m 239.463 370.731 l 236.472 357.897 m 239.463 357.897 l 237.874 370.555 m 237.874 358.073 l 224.342 380.314 m 227.333 380.314 l 224.342 370.555 m 227.333 370.555 l 226.01 380.314 m 226.01 370.731 l 213.084 409.063 m 216.074 409.063 l 213.084 388.489 m 216.074 388.489 l 214.768 408.887 m 214.768 388.489 l 201.482 410.73 m 204.472 410.73 l 201.482 396.841 m 204.472 396.841 l 202.713 410.73 m 202.713 396.841 l 189.612 423.48 m 192.602 423.48 l 189.612 399.304 m 192.602 399.304 l 191.023 423.48 m 191.023 399.304 l 154.278 341.721 m 157.268 341.721 l 154.278 346.731 m 157.268 346.731 l 155.593 341.897 m 155.593 346.906 l 274.036 366.22 m 283.8 363.574 l 239.243 364.75 m 271.537 365.632 l 227.065 374.22 m 237.27 364.456 l 215.243 397.044 m 225.301 376.572 l 203.947 403.574 m 213.564 398.955 l 192.272 410.545 m 201.301 404.603 l 189.858 410.042 m 156.829 345.601 l 154.918 343.249 m 145.154 335.543 l 134.006 334.955 m 143.243 334.808 l S BT /F18 1 Tf 10 0 0 10 118.142 319.31 Tm /GS3 gs 1.9099 Tc [(02)-17.1(4)75.3(6)-51.1(8)272.3(1)1909.9(0)575(1)1909.9(2)438.9(1)1909.9(4)]TJ -4.2857 -3.1745 TD 0 Tc (c\))Tj /F19 1 Tf 6 0 0 6 282.24 214.186 Tm (l)Tj /F18 1 Tf 0 10 -10 0 95.786 201.982 Tm (TER )Tj /F20 1 Tf 1.868 0 TD (W)Tj /F18 1 Tf 0.768 0 TD (\341cm)Tj 0 8 -8 0 92.951 241.551 Tm (2)Tj ET /GS4 gs 120.368 283.184 m 120.368 171.502 l 120.632 282.992 m 117.89 282.992 l 120.632 263.246 m 117.89 263.246 l 120.632 245.195 m 117.89 245.195 l 120.632 226.801 m 117.89 226.801 l 120.632 189.257 m 117.89 189.257 l 120.632 171.461 m 117.89 171.461 l S BT 10 0 0 10 111.673 167.722 Tm /GS3 gs (0)Tj -0.4082 11.1618 TD (60)Tj /F19 1 Tf 6 0 0 6 164.096 211.251 Tm (l)Tj 7.7847 1.4543 TD (l)Tj -3.8199 -5.6943 TD (l)Tj ET /GS4 gs 283.065 228.167 m 286.055 228.167 l 283.065 204.189 m 286.055 204.189 l 284.648 228.161 m 284.648 204.189 l 211.495 231.611 m 214.486 231.611 l 165.263 217.943 m 168.253 217.943 l 213.094 231.571 m 213.094 212.253 l S BT /F18 1 Tf 10 0 0 10 107.592 259.483 Tm /GS3 gs (50)Tj 0 -1.7672 TD (40)Tj 0 -1.8728 TD (30)Tj ET /GS4 gs 120.632 207.43 m 117.89 207.43 l S BT 10 0 0 10 107.592 203.713 Tm /GS3 gs (20)Tj 0 -1.8057 TD (10)Tj ET /GS4 gs 119.575 168.2 m 291.998 168.2 l 119.788 168.409 m 119.788 165.667 l 143.518 168.409 m 143.518 165.667 l 167.247 168.409 m 167.247 165.667 l 190.457 168.409 m 190.457 165.667 l 214.201 168.409 m 214.201 165.667 l 237.307 168.409 m 237.307 165.667 l 260.158 168.409 m 260.158 165.667 l 283.902 168.409 m 283.902 165.667 l S BT 10 0 0 10 117.575 155.913 Tm /GS3 gs 1.9099 Tc [(01)39.6(2)18.6(3)5.6(4)45.6(5)62.3(6)-17.1(7)]TJ ET /GS4 gs 211.495 211.97 m 214.486 211.97 l 188.755 180.951 m 191.746 180.951 l 188.755 193.151 m 191.746 193.151 l 190.223 193.155 m 190.223 181.234 l 165.263 209.524 m 168.253 209.524 l 166.644 217.995 m 166.644 209.715 l S BT /F19 1 Tf 6 0 0 6 141.234 260.791 Tm /GS3 gs 0 Tc (l)Tj ET /GS4 gs 142.103 248.263 m 145.094 248.263 l 142.103 277.042 m 145.094 277.042 l 143.484 277.093 m 143.484 248.454 l 143.713 262.073 m 165.713 214.574 l 189.463 188.824 m 166.463 212.574 l 212.713 221.074 m 191.213 188.824 l 283.463 216.324 m 213.963 221.574 l S BT /F18 1 Tf 10 0 0 10 173.997 144.413 Tm /GS3 gs [(Da)30(ys after plating)]TJ 13.9802 31.1653 TD (b\))Tj ET /GS4 gs 346.399 451.683 m 346.399 334.711 l 346.662 451.491 m 343.921 451.491 l 346.662 427.759 m 343.921 427.759 l 346.662 404.665 m 343.921 404.665 l 346.662 381.743 m 343.921 381.743 l 346.662 357.756 m 343.921 357.756 l 346.662 334.79 m 343.921 334.79 l S BT 10 0 0 10 337.704 331.565 Tm /GS3 gs (0)Tj ET /GS4 gs 346.173 331.597 m 515.961 331.597 l 346.385 331.806 m 346.385 329.065 l 370.115 331.806 m 370.115 329.065 l 393.845 331.806 m 393.845 329.065 l 417.054 331.806 m 417.054 329.065 l 440.799 331.806 m 440.799 329.065 l 463.905 331.806 m 463.905 329.065 l 486.755 331.806 m 486.755 329.065 l 510.5 331.806 m 510.5 329.065 l S BT 10 0 0 10 328.864 354.431 Tm /GS3 gs (100)Tj 0 2.358 TD (200)Tj 0.0102 2.3087 TD (300)Tj 0 2.3248 TD (400)Tj 0 2.3493 TD (500)Tj /F19 1 Tf 6 0 0 6 494.558 362.657 Tm (l)Tj -7.7129 6.6877 TD (l)Tj -5.7293 -4.1657 TD (l)Tj 3.8662 6.3069 TD (l)Tj -9.6314 -4.3183 TD (l)Tj -1.8819 -8.2673 TD (l)Tj -1.8819 -1.1486 TD (l)Tj ET /GS4 gs 495.44 361.885 m 498.431 361.885 l 449.218 409.103 m 452.209 409.103 l 437.871 429.333 m 440.862 429.333 l 414.554 382.682 m 417.545 382.682 l S BT 6 0 0 6 505.479 359.776 Tm /GS3 gs (l)Tj ET /GS4 gs 506.261 366.534 m 509.251 366.534 l 506.261 357.962 m 509.251 357.962 l 507.844 366.384 m 507.844 357.917 l 495.44 367.709 m 498.431 367.709 l 496.738 367.627 m 496.738 361.803 l 449.218 401.258 m 452.209 401.258 l 450.676 409.026 m 450.676 401.492 l 437.871 407.043 m 440.862 407.043 l 439.494 429.295 m 439.494 407.005 l 414.554 377.169 m 417.545 377.169 l 415.961 382.694 m 415.961 377.181 l 379.912 399.615 m 382.903 399.615 l 379.912 385.17 m 382.903 385.17 l 381.319 399.627 m 381.319 385.181 l 360.427 336.249 m 369.515 341.762 l 370.676 343.783 m 381.163 391.316 l 415.184 380.829 m 382.562 391.782 l 416.893 381.14 m 438.406 416.87 l 450.137 405.762 m 439.961 417.492 l 495.96 365.762 m 451.764 404.052 l 497.981 364.208 m 506.831 362.498 l S BT /F18 1 Tf 10 0 0 10 344.275 319.31 Tm /GS3 gs 1.91 Tc [(02)-16.9(4)75.4(6)-51(8)272.4(1)1910(0)575.1(1)1910(2)439(1)1910(4)]TJ -3.9384 -3.1812 TD 0 Tc (d\))Tj ET /GS4 gs 346.399 283.073 m 346.399 167.82 l 346.662 283.073 m 343.921 283.073 l 346.662 225.814 m 343.921 225.814 l 510.461 168.074 m 350.462 168.074 l S BT 10 0 0 10 327.486 279.134 Tm /GS3 gs (10)Tj 8 0 0 8 336.606 281.968 Tm (-5)Tj 10 0 0 10 327.486 222.247 Tm (10)Tj 8 0 0 8 336.606 225.081 Tm (-6)Tj 10 0 0 10 327.486 164.622 Tm (10)Tj 8 0 0 8 336.606 167.456 Tm (-7)Tj ET /GS4 gs 353.121 231.251 39.685 -63.177 re 407.871 247.324 39.685 -79.25 re 463.121 247.324 39.685 -79.25 re 363.212 237.323 m 383.462 237.323 l 373.462 231.574 m 373.462 237.323 l 418.212 249.323 m 438.462 249.323 l 428.462 247.324 m 428.462 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[(desm)-7.8(oplakin)-283.7(revealed)-284.8(the)-278.2(presen)-8.2(ce)-275.5(of)-276.3(a)-280(puncta)-7.8(te)-277.5(staining)]TJ 0 -1.0755 TD [(pattern)-234.4(typical)-229.5(of)-230.7(desmosom)-9.5(es)-225.1(\(fig)-8.6(s.)-224.3(2C)-228.2(and)-230.3(3C\).)-225.2(A)-226.6(maj)-8.2(o-)]TJ T* [(rity)-254.1(of)-253.5(the)-255.5(stai)-9.4(ning)-255.8(was)-252.5(associ)-10.1(ated)-256.6(with)-256.4(the)-255.5(cell)-253.3(perip)-8.2(hery)57.5(,)]TJ T* [(althou)-9.7(gh)-297.4(numer)-9(ous)-299.4(puncta)-309.4(were)-305.3(eviden)-7.8(t)-299.1(within)-305.9(the)-301(cell)-8.6(s.)]TJ T* [(In)-310.4(both)-312.7(of)-304.7(the)-312.4(human)-314.7(airway)-314.9(cell)-310.2(line)-8.2(s)-309.2(the)-306.7(pa)-7.6(tterns)-314.1(of)-304.7(E-)]TJ T* [(cadheri)-10.5(n)-347.9(and)-355.5(desmop)-9.3(lakin)-350.4(imm)-11(unostaini)-9.9(ng)-348.7(wer)-8.2(e)-348.3(simil)-9.6(ar)]TJ T* [(to)-334(those)-337.8(seen)-340.8(in)-334(MD)-7.4(CK)-334.5(cells)-340.6(\(not)-337.1(shown\).)]TJ 1.0015 -1.0471 TD [(Co)16.5(mp)18.1(ar)13.5(is)17.7(on)-269(of)-264.9(fi)17.3(gu)21.2(re)-271(2A)-198.9(wi)-7.8(th)-288.4(figur)-9.4(e)-291.4(2D,)-290.4(and)-292.9(of)-282(figur)-9.4(e)]TJ -1.0015 -1.0755 TD [(3A)-125(wi)-7.8(th)-208.8(figure)-215.4(3D,)-210.7(shows)-209.3(that)-213.1(the)-215.6(majority)-217.6(of)-208(the)-209.9(occlud)-7.8(in)]TJ T* [(in)-527.4(both)-529(16HBE14o)-7.5(-)-525.9(and)-526.2(Calu-3)-531.5(cell)-8.6(s)-525.5(appeared)-532.8(to)-527.4(be)]TJ T* [(strong)-9.7(ly)-214.5(associated)-221.2(with)-222.3(the)-215.6(cytoskelet)-11.5(on)-212.1(since)-218.7(the)-215.6(patt)-8.2(ern)]TJ T* [(of)-213.7(immunos)-9.5(taining)-212.1(was)-212.7(unc)-8.4(hanged)-210.2(by)-212.1(detergent)-207.9(treatmen)-11.1(t)]TJ T* [(of)-327.5(the)-329.4(cells)-329.2(prior)-328.1(to)-328.3(fixation.)-334.3(This)-329.4(confi)-8.2(rms)-326.9(the)-329.4(expect)-8.2(ed)]TJ T* [(strong)-368.2(associati)-11.9(on)-360(between)-371.4(functional)-370(TJs)-367.3(and)-361.2(the)-363.6(cyto-)]TJ T* 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[(\(com)-8.2(pare)-466.4(fig.)-461.9(3C)-461.5(an)-7.6(d)-456(F\).)-463.6(The)-467.8(authors)-466.2(infer)-465(that)-463.5(these)]TJ T* [(parti)-10.6(cular)-367.1(compo)-7.4(nents)-366.3(of)-361.6(inte)-8.2(rcellular)-371.2(junctions)-368.4(in)-368.1(16H-)]TJ T* [(BE)-238(14o-)-238(and)-236(Calu-3)-241.3(cells)-238.2(are)-232.3(less)-241.5(well)-234.1(associ)-10.1(ated)-239.6(with)-233.7(the)]TJ T* [(cytos)-7.8(kelet)-9.4(on)-382.8(than)-387.1(are)-386(the)-380.7(TJ)-388.1(proteins;)-392.8(the)-380.6(de)-7.6(ter)10.4(gent)-387.1(ex-)]TJ T* [(tract)-11(able)-518.4(fraction)-527(perhaps)-521.1(repres)-9.8(enting)-521.9(E-cadherin)-525(and)]TJ 0 -1.0698 TD [(desm)-7.8(oplaki)-9.7(n)-325.1(w)0(i)-7.8(t)0(h)-328.3(mai)-8.2(nly)-329(extra)-8.6(junctional)-342.3(or)-327.5(cytopl)-9.7(asmic)]TJ 0 -1.0755 TD [(local)-9.4(ization.)-565.7(Rhoda)-8.9(mine-phal)-11.5(loidin)-563.7(staining)-566.8(of)-560.8(F-actin)]TJ T* 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u,two1`3ˈUF '{k]X|EOԧX/T(ەv =W) £Mƀ@Ruэc_zsk:فastxӒY @:\=o[o?nPgT:W%3=āA<-y 2'F(I, Wnblnhx,kgJuۣ-aBF}.DIm{wɶ ~ZVxF)LXYHe*q\CA7O7mgۻ÷?=jSSqf&uolF[i.u2jo*)7UZГ۹~Z#e1=H. c;'9ǩFq h_pIK  3)e#}HlUw2I pg[G, ]2s[~P =g v|#IbĒ5g5X0gѻo<2?]ɔrPJ Z\9nqηuLQTyͶlqԮ yD۱M {I:u "+(@G/${x%z03G:*<~ )MWBopWUf+/}oϷկ?=JGX [Cbo?V) s/nR.I["Nlj;u5j"Y8яlqs_ZuK0-o Ǐ>fzR¿+EL"ܙ%EoC+g1<~Wnuһ]ĕ67}^pX .5|9l 6:`e5]ԞO%af=nߙ/ʋoӷnӻdcII s< Ԅu{iRb1 pGsPV9\^ě[%G,jYwt:fKi7^?Ը\͟d1m.6{qFQ5]lՋigҪ6 =qJUnu$Zq2~:eUO_2-w0؃pXP뇭^:WB~Nm}noճzeϹ3E@݉'5,dX|=,K=͹٤Em;_6 7mmߟӽ__o86,4[SlO p7}?.?&;[Լ_ EtV+rqmߕս HW/s>y w=:`[~W?ֽO{Nns$7ߏoi5|> stream AdobedC +   o!1AQ"aq2 #BR3$bC4% rS&cD5Ts6'E7F򃓣dU()8GHVe*9:IJWXYZftugvwhijxyz? <߾}*o<{'wP~ 4[n}>мꙇH?#HŬyoe-;~w?Ѽv@(ksJȧn|-~k) kһcm׹Zݸ3=ӱoe~plG A;n0.* Er='T2N00bmԒokm[/[u]/o2?g♌dΟfs}d0j%7U38VR=挕'?jI;$t"׷ǚDq1N7B5cA7c4!^qTfF8…'A}@Z߯=9;: lŋ˘z)"o{-r_]GmCaGovxyI gөԥ&±,ًb~jY6%o}+N13]^G/LN*I PB{ߟ!ឞꙪdk6ȾX۱*~=zvX RX S3FA7h I/'??ղ H!7a# \YY79ZM: b Aͽj)b%l2u$ܑbovlM>"0LRKH藧Ʉ+!,RYۖ=O+Iַv=[sָ^8~'T@LG >MmT*M&W&f;*a$DPY 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[(ex)15.1(pl)18.9(ai)18.4(ne)15.1(d)-353.6(b)0(y)-348.7(t)0(h)13.2(e)-359.7(ag)20.8(gr)19.6(eg)15.1(at)18.4(e)-359.7(b)0(r)19.6(e)0(a)14.7(k)-359.3(le)18.4(ng)15.5(t)13.9(h)-359.3(be)15.1(in)18.9(g)-359.3(s)0(i)17.7(m)0(i)15.7(l)0(a)18.4(r)-360.9(in)]TJ 0 -1.0471 TD [(th)18.9(e)-274.3(t)0(w)20.6(o)-273.9(ce)20.4(ll)-262.4(li)16.5(ne)15.1(s;)-261.2(th)18.9(e)-274.3(g)0(a)20.8(p)0(s)-264.5(i)0(n)-260(t)0(h)18.9(e)-274.3(T)12.2(J)0(s)-265.7(o)0(f)-259.2(C)0(a)16.1(l)0(u)18.9(-)0(3)-264.9(c)0(e)20.4(l)0(l)16.5(s)-275.1(w)12.4(e)0(r)13.5(e)]TJ 0 -1.0528 TD [(nu)15.5(me)17.7(ro)19.6(us)-440.9(an)15.1(d)-450.3(o)0(f)-441.3(s)0(h)20(o)0(r)19.6(t)-452.7(di)18.9(st)17.7(an)15.1(ce)20.4(,)-449.9(w)0(h)17.3(e)0(r)13.5(e)0(a)20.4(s)-451.5(ga)15.1(ps)-440.9(in)-436.4(16)15.5(H-)]TJ 0 -1.0471 TD 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[(possib)-9.3(le)-442.5(for)-449.3(some)-451.7(TJ)-445(adhesi)-9(on)-445.4(prote)-8.2(ins)-444(to)-447.8(have)-447.7(min)-7.8(or)]TJ 25.1977 66.1583 TD [(disco)-7.8(ntinu)-8.6(ities)-484.9(in)-487.6(their)-488.6(staining)-492.9(pattern)-490.5(without)-494.4(barrier)]TJ 0 -1.0471 TD [(funct)-8.2(ion)-454.2(being)-456.2(impaired)-10.7(.)-449.9(This)-454.6(unders)-7.8(cores)-451.3(the)-454.6(need)-453.4(to)]TJ 0 -1.0528 TD [(compl)-9.7(ement)-243.5(morp)-7.8(hologi)-9.3(cal)-238.8(techni)-10.1(ques)-243.7(with)-245.1(tests)-244.7(of)-230.8(func-)]TJ T* [(tion)-389.1(when)-388.7(exploring)-389.2(the)-386.3(role)-388.7(of)-384.4(the)-386.3(epith)-9(elial)-387.3(barrier)-388.9(in)]TJ 0 -1.0471 TD [(pathop)-7.4(hysio)-7.4(logical)-343.3(events.)]TJ 1.0015 -1.0527 TD [(Im)10.8(mu)18.1(no)21.2(bl)13.2(ot)18.9(ti)16.5(ng)-229.1(de)15.1(mo)18.1(ns)20(tr)17.3(at)18.5(ed)-235.2(th)18.9(at)-231.9(th)18.8(e)-245.9(T)12.2(J)-241(p)0(r)14(o)0(t)18.9(e)0(i)18.5(n)-239.8(oc)15.1(cl)18.5(u-)]TJ -1.0015 -1.0471 TD [(di)13.2(n)-285.3(w)0(a)16.9(s)-286.5(de)20.8(te)12.8(ct)18.5(ab)20.8(le)-277.5(at)-277.5(a)-285.7(r)0(a)13.5(n)0(g)21.2(e)-285.7(of)-282(mo)18.1(le)18.4(cu)15.1(la)18.4(r)-286.9(m)0(a)17.7(s)0(s)18.8(e)0(s)-282(i)13.9(n)-291(b)0(o)21.2(t)0(h)]TJ 0 -1.0527 TD [(ce)14.7(ll)-239.6(li)16.5(ne)20.8(s)-246.7(f)0(r)18(o)0(m)-238(h)0(u)15.5(m)0(a)17.7(n)-245.5(ai)18.4(rw)15.7(ay)15.1(s)-246.7(e)0(p)20.8(i)0(t)16.5(h)0(e)15.1(l)0(i)16.5(u)0(m)18.1(.)-245.1(Th)17.1(e)-245.9(m)0(o)18.1(s)0(t)-238.4(a)0(b)15.1(u)0(n)21.2(-)]TJ 0 -1.0471 TD [(da)15.1(nt)-197.4(of)-196.6(th)18.8(e)-206(f)0(o)13.9(r)0(m)16.5(s)-201.1(ha)15.1(d)-205.6(a)0(n)-195.4(a)0(p)15.1(p)0(a)20.8(r)0(e)13.5(n)0(t)-197.4(m)0(o)18.1(l)0(e)18.4(c)0(u)15.1(l)0(a)18.4(r)-207.2(ma)17.7(ss)-197.4(of)-196.6(62)21.2(\2616)15.5(6)]TJ 0 -1.0527 TD 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[(fo)13.9(rm)16.5(s)-326.3(o)0(f)-321.8(o)0(c)15.1(c)0(l)18.5(u)0(d)15.5(i)0(n)-316.9(t)0(h)13.2(a)0(t)-317.3(s)0(e)14(a)0(l)-317.3(T)0(J)15.9(s)-332([3)19.6(3,)-331.2(34].)-340(The)-342.6(human)-343.1(air-)]TJ 0 -1.0471 TD [(way)-302.5(cell)-298.8(lines)-300.4(expres)-8.2(sed)-294.1(heav)-9.6(y)-291(form)-9.4(s)-292.2(o)0(f)-299(occlud)-7.8(in)-294.1(\(bands)]TJ 0 -1.0527 TD [(of)-338.9(~72\26174)-337.8(kDa)-331(wer)-8.2(e)-331.2(detect)-10.6(able)-336.3(in)-334(both)-335.5(cell)-333(line)-8.2(s)-332(with)-336.1(a)]TJ T* [(prote)-8.2(in)-385.2(of)-384.4(94)-382.8(kDa)-387.9(additional)-12.3(ly)-379.5(presen)-8.2(t)-384.4(i)0(n)-385.2(Calu-3)-389.2(cells\),)]TJ 0 -1.0471 TD [(alth)-8.2(ough)-509.5(the)-511.5(author)-9(s)-508.4(are)-511.2(unable)-508.5(to)-510.4(comm)-9.7(ent)-511.5(on)-508(their)]TJ 0 -1.0527 TD [(pho)-8(sphorylation)-429.2(stat)-9.4(us.)-418.5(W)72(e)-6.9(ak)-417.3(bands)-426.5(at)-425.4(43\26145)-424.9(kDa)-422(and)]TJ 0 -1.0471 TD [(52\261)-8(57)-320.2(kDa)-325.3(found)-324.1(in)-322.6(occlud)-7.8(in)-322.6(immunob)-8.3(lots)-327.7(may)-324.5(be)-320.6(pro-)]TJ 0 -1.0527 TD [(teol)-8.2(ytic)-435(fragmen)-12.5(ts)-431.9(formed)-438.1(durin)-7.8(g)-427.6(prote)-8.2(in)-430.7(extra)-8.6(ction.)-436.1(In)]TJ 0 -1.0471 TD [(con)-8.4(trast,)-461.7(occlud)-7.8(in)-459.2(from)-464.6(MD)-7.4(CK)-459.7(cell)-8.6(s)-457.2(show)-8.2(ed)-457.2(less)-463.5(het-)]TJ 0 -1.0527 TD [(erogene)-8.2(ity)-325.7(than)-318.8(its)-321.2(human)-326(counterpart)-11.9(s.)-315.3(How)-10.2(ever)28.7(,)-319.1(a)0(s)-316.1(the)]TJ 0 -1.0471 TD [(author)-9(s)-286.5(were)-288.2(not)-289.2(able)-285.1(to)-288.4(blot)-286.7(canine)-292.7(and)-287.2(human)-291.9(occludin)]TJ 0 -1.0527 TD [(wi)-7.8(th)-220.2(the)-227(same)-224.4(anti)-8.2(body)-225(because)-224.4(of)-225.1(a)-223.1(lack)-228.2(of)-219.4(speci)-8.2(es)-225.1(cross-)]TJ 0 -1.0471 TD [(react)-9(ivity)57.3(,)-290.6(cauti)-9.4(on)-291.8(should)-292.7(be)-292.2(exerci)-9.8(sed)-294.1(in)-294.1(interpret)-12.3(ing)-289.2(the)]TJ 0 -1.0528 TD [(possi)-8.6(ble)-392(meaning)-398.6(of)-390.1(this)-390.3(difference.)-392.4(All)-392.2(three)-395.6(cell)-395.6(lines)]TJ T* [(exp)-8.4(ressed)-521.5(broad)-518(b)-6.5(ands)-516.8(of)-521(ZO-1)-519.2(immunos)-9.5(tain)-8.2(ing)-516.8(with)]TJ 0 -1.0471 TD [(app)-8.4(arent)-219.2(molecular)-228.1(masses)-222.6(of)-213.7(~220)-222.4(kDa.)-217.6(The)-217.4(broad)-222.1(bands)]TJ 0 -1.0528 TD [(of)-412.8(ZO-1)-416.7(in)-407.9(immuno)-13.2(blots)-413.8(of)-407.2(all)-417.2(three)-412.6(cell)-412.7(types)-417.5(is)-409.1(con-)]TJ 0 -1.0471 TD [(siste)-11.4(nt)-499(wi)-7.8(th)-504.7(some)-502.9(heter)-8.6(ogenei)-7.8(ty)-504.7(in)-504.7(this)-504.1(prote)-8.2(in,)-505.1(either)]TJ 0 -1.0528 TD [(bec)-8.8(ause)-540(of)-538(differential)-542.5(phospho)-6.5(rylation)-544.5(or)-538(due)-537.6(to)-538.8(the)]TJ 0 -1.0471 TD 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-1.1128 TD [(19.)-1266.5(W)32.8(inton)-419.5(BL,)-421.1(W)77.1(a)0(n)-422.4(H)0(,)-416.7(Cannell)-422.8(MB,)]TJ /F4 1 Tf 17.1477 0 TD [(et)-418.5(al)]TJ /F3 1 Tf 1.8969 0 TD [(.)-413.9(Class)-424(specific)]TJ -16.5281 -1.1128 TD [(inhibition)-436.8(of)-429.9(house)-434.7(dust)-431.5(mite)-434.3(proteinases)-431.8(which)-432.9(cleave)]TJ 0 -1.1191 TD [(cell)-394.3(adhesion,)-387.6(induce)-390.9(cell)-394.3(death)-385.1(and)-390.3(which)-394.9(increase)-391.6(the)]TJ 0 -1.1128 TD [(permeability)-384.9(of)-379.3(lung)-373.4(epithelium.)]TJ /F4 1 Tf 13.569 0 TD [(Br)-383(J)-378.7(Pharmacol)]TJ /F3 1 Tf 6.9805 0 TD (1998;)Tj -20.5495 -1.1192 TD [(124:)-335.5(1048\2611059.)]TJ -2.5165 -1.1128 TD [(20.)-1266.5(Saltou)-316.2(M,)-319.2(Ando-Akatsuka)-320.2(Y)123.6(,)-312.7(Itoh)-317.5(M,)]TJ /F4 1 Tf 17.9444 0 TD [(et)-317.3(al)]TJ /F3 1 Tf 1.7957 0 TD [(.)-312.7(Mammalian,)]TJ -17.2236 -1.1128 TD 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