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[(challenges)-384.4(to)-370.9(be)-377.7(addressed,)-379.3(including)-374.3(limited)-379.2(maturity)-380.2(and)]TJ 0 -1.1116 TD [(size,)-482.4(lack)-484.9(of)-474.5(vascularization,)-486.8(and)-482.5(inability)-478.3(to)-480.4(recapitulate)]TJ T* [(communication)-516(between)-519.1(organs,)-512.1(it)-509.3(is)-509(still)-512.7(a)-509.1(good)-511.3(model)]TJ 0 -1.1174 TD [(system)-527.1(for)-527.4(regenerative)-531.8(medicine)-526.4(research.)]TJ 7.3823 0 0 6.5614 493.1715 532.2329 Tm (23,24)Tj 9.843 0 0 9.843 514.9416 527.8676 Tm (Organoid)Tj -20.5103 -1.1116 TD [(opens)-363.6(up)-365.1(a)-359.4(new)-362.4(avenue)-361.4(for)-366.1(pathogenesis)-362.2(research)-370.3(and)-361.6(per-)]TJ T* [(sonali)-7.8(zed)-506.7(drug)-511.2(screening)-514.4(for)-504.3(treatment)-516.1(of)-509(infectious)-514.6(dis-)]TJ 0 -1.1174 TD (eases,)Tj 7.3823 0 0 6.5614 336.3589 499.4077 Tm (25,26)Tj 9.843 0 0 9.843 356.2582 494.9857 Tm [(genetic)-328.2(diseases,)]TJ 7.3823 0 0 6.5614 422.8156 499.4077 Tm [(27,2)-8.6(8)]TJ 9.843 0 0 9.843 442.7148 494.9857 Tm [(and)-327(cancer.)]TJ 7.3823 0 0 6.5614 488.466 499.4077 Tm [(29,3)-8.6(0)]TJ 9.843 0 0 9.843 508.3652 494.9857 Tm [(In)-324.7(addition)]TJ -19.8422 -1.1116 TD [(to)-284.6(drug)-286.6(screening)-289.8(and)-280.9(molecular)-291.3(medicine,)]TJ 7.3823 0 0 6.5614 483.1936 488.4093 Tm [(25,2)-8.6(9)]TJ 9.843 0 0 9.843 502.6392 484.0439 Tm [(the)-286.8(potential)]TJ -19.2605 -1.1116 TD [(value)-624.3(of)-624.2(organoid)-620.6(systems)-620.2(in)-624.4(basic)-618.3(biological)-624.4(research)]TJ 0 -1.1174 TD [(should)-306.1(not)-302.9(be)-302.9(understated.)]TJ 7.3823 0 0 6.5614 418.1668 466.5258 Tm (31)Tj 9.843 0 0 9.843 428.5416 462.1038 Tm [(Studies)-302.5(on)-301.7(optic)-306.4(cup)-298.2(organoids)]TJ -11.7325 -1.1116 TD [(have)-306.2(revealed)-301.5(the)-304.1(morphogenetic)-307.3(mechanisms)-304.7(of)-295.9(optic)-306.4(cups)]TJ T* [(and)-263.7(their)-260.3(species)-264.4(specicity.)]TJ 7.3823 0 0 6.5614 424.7432 444.5857 Tm (32,33)Tj 9.843 0 0 9.843 443.9054 440.2204 Tm [(In)-261.4(addition,)-261.5(organoids)-264.4(have)]TJ -13.2934 -1.1116 TD [(demonstrated)-386.6(their)-381.3(potential)-375.6(as)-383.3(a)-376.6(minimalist)-383.7(model)-376.7(system)]TJ 0 -1.1174 TD [(to)-699.2(study)-702.4(the)-701.5(principle)-703.8(of)-699.1(self-organization.)]TJ 7.3823 0 0 6.5614 501.392 422.6456 Tm (34)Tj /F7 1 Tf 9.843 0 0 9.843 515.6786 418.2802 Tm [(In)-699.1(silico)]TJ /F6 1 Tf -20.5852 -1.1116 TD [(mathematical)-431(modeling)-426.2(has)-419(also)-426(been)-421.4(used)-425.8(to)-422.8(explore)-424.8(the)]TJ T* [(molecular)-619.6(mechanism)-609(of)-612.7(organoid)-614.8(self-organization)-616.9(and)]TJ 0 -1.1174 TD [(predict)-240.7(the)-240.7(impact)-240.8(of)-238.3(various)-239.2(microenvironmental)-246.3(conditions)]TJ 0 -1.1116 TD [(on)-336.3(culture)-338.6(results.)]TJ 7.3823 0 0 6.5614 385.1716 378.8219 Tm (35,36)Tj /F9 1 Tf 8.9663 0 0 8.9663 313.0582 346.9605 Tm [(Organoid)-337.8(Systems)-334.1(Are)-336.7(Distinguished)]TJ 0 -1.2203 TD [(by)-338.9(Sustainability)-335.4(in)-331.6(Long-Term)-339.2(Culture)]TJ T* [(and)-338.2(Capacity)-335.3(for)-330.9(Self-Organization)]TJ /F6 1 Tf 9.843 0 0 9.843 323.0361 308.6361 Tm [(Organoid)-471.7(technology)-462.9(debuted)-470.9(more)-467.5(than)-466.5(10)-463(years)-468.4(ago,)]TJ -1.0137 -1.1116 TD [(and)-321.2(since)-318.8(then,)-320.2(various)-319.8(organoid)-321.1(culture)-321.3(systems)-320.7(have)-317.8(been)]TJ 0 -1.1174 TD [(devel)-8(oped.)-366.1(In)-365(addition)-373.2(to)-365.2(extracellular)-371.9(matrix)-372.1(\(ECM\))-371.4(scaf-)]TJ 0 -1.1116 TD [(folds,)-400.5(organoid)-390.2(suspension)-397.6(culture)-396.2(systems,)-399(rotating)-396.2(biore-)]TJ T* [(actor,)-328.2(air-liquid)-329.4(interface)-325.8(\(ALI\))-330(method,)-323.6(and)-327(other)-323.6(methods)]TJ 0 -1.1174 TD [(can)-426.1(also)-426(be)-423.8(used)-420.1(to)-422.8(produce)-424.7(organoids.)]TJ 7.3823 0 0 6.5614 475.8235 258.1795 Tm (13)Tj 9.843 0 0 9.843 487.3889 253.8141 Tm [(No)-423.5(matter)-425.1(what)]TJ -17.7111 -1.1116 TD [(method)-504.5(is)-497.5(used,)-498.4(to)-503.4(label)-497.7(any)-499.8(3D)-504.2(culture)-499.8(as)-498.5(an)-498.7(organoid)]TJ T* [(system,)-611.2(the)-609.4(following)-615.9(two)-609.1(core)-605.7(chara)-9(cteristics)-613.6(must)-604.6(be)]TJ 0 -1.1174 TD (present:)Tj 1.0137 -1.1116 TD [(First,)-417.7(stem)-415.7(cells)-416.8(are)-420.3(able)-415.8(to)-417(self-renew)-418.4(and)-419.2(mature)-419.2(into)]TJ -1.0137 -1.1116 TD [(terminally)-262.7(differentiated)-262.4(cells)-255.6(under)-260.1(articial)]TJ /F7 1 Tf 18.3159 0 TD [(in)-255.8(vitro)]TJ /F6 1 Tf 3.1851 0 TD (culture)Tj -21.501 -1.1174 TD [(conditions.)-246.2(The)-247.6(self-renewal)-249.1(capacity)-244.1(allows)-244.9(organoids)-247.1(to)-238.5(be)]TJ 0 -1.1116 TD [(cultured)-391.5(for)-389.1(up)-382.3(to)-388.2(more)-386.9(than)-391.6(a)-382.4(year,)-390.3(which)-387.8(contain)-390.4(prolif-)]TJ T* [(erating)-563.2(cells)-560.8(throughout)-558.4(the)-557.5(entire)-562.1(culture)-563.2(period,)-558.5(while)]TJ T* [(plurip)-6.8(otency)-354.6(allows)-360.1(stem)-358.1(cells)-353.5(to)-359.4(differentiate)-359.2(and)-355.8(produce)]TJ 0 -1.1174 TD [(different)-375.3(cell)-369.9(lineages,)-367.1(similar)-373.1(to)-365.2(normal)-366.2(tissues.)]TJ 7.3823 0 0 6.5614 509.8959 148.5921 Tm (37)Tj 9.843 0 0 9.843 520.8943 144.2267 Tm [(It)-365.2(must,)]TJ -21.1151 -1.1116 TD [(howev)-7.5(er,)-657.6(be)-660(noted)-663.4(that)-662.4(by)-658.8(denition,)-665.6(the)-661.2(self-renewal)]TJ T* [(property)-407.5(should)-404(not)-400.9(be)-400.8(attributed)-405.5(to)-405.5(organoids,)-406.1(but)-400.9(should)]TJ 0 -1.1174 TD [(instead)-401.8(be)-389.3(assigned)-396.8(to)-394(stem)-398.4(cells.)-397.2(In)-393.8(the)-390.5(organoid)-396(passage)]TJ 0 -1.1116 TD [(process,)-324.1(an)-320.1(organoid)-321.1(is)-318.9(dissociated)-320.9(into)-321.4(multiple)-317(fragments.)]TJ T* [(Some)-381(fragments)-382(contain)-384.7(stem)-381.1(cells,)-379.9(which)-382.1(then)-380.1(form)-379.9(mul-)]TJ 0 -1.1174 TD [(tiple)-669.4(organoids,)-665.3(creating)-669.1(the)-661.2(illusion)-665.7(of)-664.5(organoid)-666.7(self-)]TJ 0 -1.1116 TD [(renewal.)-522.5(Rationally,)-513.7(the)-517.2(ability)-516.2(of)-509(organoids)-517.8(to)-514.9(replicate)]TJ T* [(thems)-7.8(elves)-347.6(and)-344.3(their)-352.5(sustainability)-346.4(in)-347.9(long-term)-352.3(culture)-350.1(are)]TJ /F9 1 Tf 8.9663 0 0 8.9663 62.022 749.6502 Tm (546)Tj 48.541 0 TD [(QING)-334.9(ET)-338.7(AL.)]TJ ET endstream endobj 54 0 obj <>stream /GS1 gs BT /F6 1 Tf 9.843 0 0 9.843 59.7543 374.57 Tm 0 0 0 rg 0 Tc 0 Tw [(due)-246.4(to)-250(the)-246.5(self-renewal)-249.1(capacity)-249.9(of)-244.1(stem)-248.6(cells)-249.8(existing)-246.3(within)]TJ 0 -1.1116 TD [(organoid)-338.4(fragments.)]TJ 1.0137 -1.1174 TD 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[(ura)12.8(l)-329.5(bon)14(e,)]TJ 7.3823 0 0 6.5614 99.6094 115.7102 Tm [(41)17.8(43)]TJ 9.843 0 0 9.843 121.096 111.3448 Tm [(car)11.7(til)13.6(age)11.7(,)]TJ 7.3823 0 0 6.5614 157.2094 115.7102 Tm (44)Tj 9.843 0 0 9.843 167.811 111.3448 Tm [(or)-318.9(bon)14(e)-330.6(m)0(a)12.7(rro)14(w)]TJ 7.3823 0 0 6.5614 231.7606 115.7102 Tm (45)Tj 9.843 0 0 9.843 242.3055 111.3448 Tm [(wit)12.8(h)-329.4(com)17.3(ple)12.7(te)]TJ -18.5463 -1.1116 TD [(fun)14(cti)12.6(ons)14.2(.)-274.1(The)-264.8(app)12.8(ear)11.7(anc)11.7(e)-273(o)0(f)-267.1(the)12.7(se)-273.9(3D)-268(mod)12.7(els)-263.6(pro)14(vid)13.8(es)-268.1(cue)11.7(s)]TJ T* [(for)-360.3(the)-361.7(con)12.8(str)14.1(uct)18.4(ion)-360.5(of)-365(ske)13(let)12.6(al)-360.6(syst)18.9(em)-367.4(org)14(ano)12.8(ids)-360.3(an)13.9(d)-369.7(hel)12.7(ps)]TJ 0 -1.1174 TD [(sha)13(pe)-279.8(thi)13.7(s)-283.1(el)12.8(d.)-275.2(How)12.2(eve)11.7(r,)-275.2(as)-279.6(exp)12.8(lai)12.6(ned)-275.2(ab)13.9(ove)12.8(,)-285.7(org)14(an)13.9(oid)13.9(s)-283.1(are)]TJ /F7 1 Tf 0 -1.1116 TD [(in)-227(vit)12.6(ro)]TJ /F6 1 Tf 3.1102 0 TD [(3D)-227.7(mod)12.7(el)-222.4(sys)14.4(tem)11.5(s,)-223.2(but)-222.3(the)-217.7(rev)12.9(ers)13(e)-232.6(sta)12.9(te)13.8(men)11.6(t)-231.6(doe)18.6(s)-231.3(not)]TJ -3.1102 -1.1116 TD [(app)12.8(ly.)-351.3(In)-359.3(thi)13.7(s)-363.8(rev)12.8(iew)11.8(,)-360.5(3)0(D)-360.2(b)0(o)15.1(n)0(e)-360.5(for)14(mat)11.5(ion)-349(mod)12.7(els)-355.7(tha)12.7(t)-358.3(wer)12(e)]TJ 25.5097 32.3062 TD [(lab)12.7(el)13.8(ed)-279.8(as)-273.9(org)14(ano)12.8(id)15(,)-279.9(inc)12.7(lud)13.8(ing)-268.4(bon)14(e)-278.7(org)14(ano)12.8(ids)14(,)-279.9(bon)14(e)-278.7(mar)11.6(ro)15.1(w)]TJ 0 -1.1116 TD [(org)14(ano)18.6(ids)14(,)-285.7(ost)14(eoc)11.7(hon)14(dra)12.8(l)-283.5(org)14(ano)12.8(ids)14(,)-285.7(c)0(a)12.8(rti)13.8(lag)12.7(ino)13.9(us)-278.5(org)14(ano)12.8(ids)14(,)]TJ 0 -1.1174 TD [(and)-465.2(cal)11.5(lus)-464(org)14(ano)12.8(id,)-466.5(wer)12(e)-474.5(car)11.7(efu)12.8(lly)-470.1(ev)13.9(alu)12.7(ate)11.5(d)-473.4(and)-471(col)12.7(le)13.8(c-)]TJ 0 -1.1116 TD [(tiv)13.7(el)13.8(y)-329.4(ref)12.9(err)12.9(ed)-325.9(to)-319.1(as)-325.7(ske)13(let)12.6(al)-326(sys)14.4(te)13.8(m)-330.6(org)14(ano)12.8(ids)14(.)]TJ 1.0137 -1.1116 TD [(Since)-330.3(2003,)-330.4(bone)-328.1(marrow)-332.3(stem)-335(cell)-329.5(\(BMSC\))-331.9(precipitates)]TJ -1.0137 -1.1116 TD [(obtained)-414.6(by)-405.4(centrifugation)-416.8(were)-416.5(induced)-413.3(to)-405.5(form)-414.5(cartilage)]TJ 0 -1.1174 TD [(and)-373.1(subsequently)-378.3(undergo)-377.5(mineralization,)-382.5(with)-374.2(the)-379(derived)]TJ 0 -1.1116 TD [(culture)-643.8(being)-640.4(known)-639.9(as)-642.5()50.8(chondro-osseous)-647.1(organoid.)48.8()]TJ 7.3823 0 0 6.5614 542.551 302.2298 Tm (46)Tj 9.843 0 0 9.843 310.8472 286.9228 Tm [(Pievani)-326.9(et)-331.8(al.)-329.5(used)-327.9()50.8(heterotopic)-333.8(bone/marrow)-333.4(organoid)46.5()-329.4(t)0(o)]TJ 0 -1.1174 TD [(refer)-249.6(to)-250(ossicles)-252.7(formed)-256.6(by)-249.9(BMSCs)-250(transplanted)]TJ /F7 1 Tf 19.4966 0 TD [(in)-250(vivo)]TJ /F6 1 Tf 2.7071 0 TD (.)Tj 7.3823 0 0 6.5614 531.8928 280.3464 Tm (47)Tj 9.843 0 0 9.843 541.7573 275.9243 Tm (In)Tj -23.4593 -1.1116 TD [(the)-551.8(past)-558.4(decade,)-556.1(various)-556(skeletal)-553.9(system)-555.9(organoids)-558.2(with)]TJ T* [(varying)-331.5(levels)-337.3(of)-336.2(complexity)-337.5(have)-335(been)-335(reported.)]TJ 1.0137 -1.1174 TD [(Ab)14.2(rah)12.9(am)-235(et)-228.1(al.)-225.8(har)12.8(ves)13(ted)-223.5(cel)11.5(l)-237.4(iso)14(lat)12.6(es)-227.8(fro)14(m)-238.5(b)0(o)15.1(ne,)-225.7(car)11.7(til)13.6(age)11.7(,)]TJ -1.0137 -1.1116 TD [(and)-476.8(rib)-481.4(tis)13.9(sue)13(s)-490.5(and)-476.8(mix)12.6(ed)-487.2(the)12.7(m)-486.1(wit)12.8(h)-490.7(end)12.8(ot)14.9(hel)12.7(ial)-482.8(cel)11.5(ls)-480.2(to)]TJ T* [(con)12.8(str)14.1(uc)13.9(t)-329.5(bon)14(e,)-328.2(ca)12.8(rti)13.7(lag)12.7(e,)-328.2(an)13.9(d)-329.4(bon)14(e-c)11.7(art)12.7(ila)12.6(ge)-325.9(or)15.1(gan)12.8(oid)13.9(s.)-326.9(Th)13.9(e)]TJ T* [(pro)14(duc)18.6(t)-243.1(cou)12.8(ld)-238.5(be)-233.7(sus)14.4(tai)12.6(nab)12.8(ly)-238.5(cu)13.9(ltu)13.7(red)]TJ /F7 1 Tf 15.1077 0 TD [(in)-238.5(vi)13.8(tro)]TJ /F6 1 Tf 3.1333 0 TD [(for)-233.6(4)-243(m)0(o)13.8(nth)13.8(s.)]TJ 7.3823 0 0 6.5614 542.6077 214.5826 Tm (48)Tj 9.843 0 0 9.843 310.8472 199.2188 Tm [(Sim)18.5(ila)12.6(rly)13.9(,)-423.9(pri)13.9(mar)11.6(y)-421.6(hum)12.7(an)-418.1(BM)13.1(SCs)-418.7(we)13(re)-423.8(ino)13.8(cu)13.9(lat)12.6(ed)-418.1(on)-422.7(po)15.1(-)]TJ 0 -1.1116 TD [(rou)14(s)-300.4(3)0(D)-291.1(sil)13.9(k)-300.6(br)14.2(oin)-291.4(sca)11.8(ffo)19.8(lds)-291.2(and)-292.4(cul)12.7(tur)13.9(ed)-291.3(in)-296.1(a)-301.8(spi)14(nne)12.8(r-)14.2(as)14.1(k)]TJ T* [(bio)13.9(rea)17.4(cto)12.7(r)-456.1(t)0(o)-457.3(o)0(b)15.1(tai)12.6(n)-456.1(wov)13.1(en)-452.6(bon)14(e)-457.3(org)14(ano)12.8(ids)14(.)]TJ 7.3823 0 0 6.5614 490.9605 181.7007 Tm (49)Tj 9.843 0 0 9.843 502.8093 177.3354 Tm [(In)-451.4(add)12.8(iti)13.6(on,)]TJ -19.5024 -1.1174 TD [(Bla)11.7(ch)13.9(e)-422.7(e)0(t)-412.4(a)0(l)13.8(.)-418.1(co-)12.8(cul)12.7(tur)13.9(ed)-412.3(mes)11.8(enc)17.4(hym)12.8(al)-412.4(ste)12.9(m)-417(cel)11.5(ls)-411.1(\()-81.7(M)0(S)14.3(Cs\))]TJ 0 -1.1116 TD [(wit)12.8(h)-294.8(end)12.8(oth)19.6(eli)12.6(al)-291.5(cel)11.5(ls)-290.1(in)-296.1(rou)14(nd)15.1(-bo)14(tto)13.7(m)-296.1(pol)13.8(y\(e)12.8(thy)13.9(len)12.7(e)-296(gly)13.8(co)13.9(l\))]TJ T* [(\(PE)13(G\))-383.2(hyd)14(rog)14(el)-383.6(mic)11.5(row)13.1(el)13.8(ls)-388(to)-382.5(pro)14(duc)12.8(e)-388.2(bon)14(e)-393.9(m)0(a)12.7(rro)14(w)-393.6(org)14(a-)]TJ 0 -1.1174 TD [(noi)13.8(ds)-226.6(wit)12.8(h)]TJ /F7 1 Tf 4.3774 0 TD [(in)-227(vi)13.8(vo)]TJ /F6 1 Tf 2.8971 0 TD [(fun)14(cti)12.6(ona)12.8(l)-231.6(cha)11.7(ra)13.9(cte)11.5(ris)14.1(tic)12.6(s.)]TJ 7.3823 0 0 6.5614 482.6833 137.8771 Tm (50)Tj 9.843 0 0 9.843 492.2644 133.455 Tm [(Hem)10.7(ato)12.7(poi)13.8(eti)12.6(c)]TJ -18.4311 -1.1116 TD [(st)20.9(em)12.7(/p)20.7(ro)15.1(ge)19.7(ni)14.9(to)20.7(r)-271.8(c)0(e)18.5(l)0(l)14.8(s)-271.6(co)19.7(ul)14.9(d)-271.8(b)0(e)-262.6(r)0(e)19.7(c)0(r)14(u)0(i)20.7(t)0(e)13.8(d)-266(an)13.9(d)-271.8(l)0(o)20.7(c)0(a)12.8(l)0(i)20.6(z)0(e)18.5(d)-271.8(ar)14(ou)20.8(nd)]TJ T* [(th)20.7(e)-238.4(e)0(n)13.9(d)0(o)20.8(t)0(h)15(e)0(l)19.5(i)0(a)13.8(l)-237.4(ne)19.7(tw)14.1(or)20.9(k.)-229.2(Al)14.1(th)20.7(ou)15.1(gh)-226.8(th)20.7(e)-238.4(l)0(o)14.9(n)0(g)20.8(-)0(t)15(e)0(r)19.7(m)-238.5(ma)12.7(in)20.7(te)13.8(na)19.7(nc)13.9(e)]TJ 0 -1.1174 TD [(of)-301.7(bo)20.8(ne)-302.9(ma)12.7(rr)20.9(ow)-302.6(or)15.1(ga)19.7(no)15.1(id)20.7(s)-311.9(w)0(a)18.8(s)-311.9(no)15.1(t)-306.5(v)0(e)13.9(r)0(i)20.7()0(e)14.1(d)0(,)-298.3(t)0(h)15(e)-313.3(se)19.9(lf)15(-r)20.9(en)13.9(ew)18.8(al)]TJ 0 -1.1116 TD [(ca)18.5(pa)13.9(ci)19.5(ty)-301.8(of)-307.4(th)20.7(e)-313.3(m)0(e)18.4(s)0(e)14.1(n)0(c)19.7(h)0(y)15.1(m)0(a)18.4(l)-312.3(co)19.7(mp)13.8(ar)19.7(tm)13.7(en)19.7(t)-312.2(w)0(a)18.8(s)-311.9(in)15(di)20.7(ca)12.8(te)19.6(d)-312.1(b)0(y)]TJ T* [(it)20.6(s)-323.4(i)0(m)13.7(m)0(u)19.6(n)0(o)15.1(p)0(h)20.8(e)0(n)13.9(o)0(t)20.7(y)0(p)15.1(e)-324.8(ma)18.4(in)20.7(te)13.8(na)19.7(nc)13.9(e.)]TJ 1.0137 -1.1116 TD [(Hu)20(ma)18.4(n)-479.1(p)0(e)13.9(r)0(i)20.7(o)0(s)15.3(t)0(e)19.5(u)0(m)13.9(-)0(d)20.9(e)0(r)14(i)0(v)20.7(e)0(d)-469.9(c)0(e)12.8(l)0(l)20.6(s)-479(\(h)15.1(PD)20.1(Cs)14.2(\))-479.1(w)0(e)18.8(r)0(e)-469.9(a)0(l)13.8(s)0(o)-468.5(r)0(e)19.7(-)]TJ -1.0137 -1.1174 TD [(po)20.8(rt)15(ed)-222.2(to)-221.2(fo)15.1(ll)20.6(ow)-227.7(th)20.7(e)-232.7(e)0(a)12.8(r)0(l)20.7(y)-231.5(pa)13.9(tt)20.6(er)14(n)-231.5(o)0(f)-221(e)0(n)13.9(d)0(o)20.8(c)0(h)13.9(o)0(n)20.8(d)0(r)15.1(a)0(l)-222.4(o)0(s)21(s)0(i)15.1()0(c)14.1(a)0(t)19.5(i)0(o)20.7(n)]TJ ET q 1 i 59.754 732.245 m 550.091 732.245 l 550.091 435.515 l 59.754 435.515 l W n q 490.48 0 0 296.731 59.754 435.515 cm /Im2 Do Q Q BT /F8 1 Tf 9.845 0 0 9.845 59.7543 418.7337 Tm [(FIG.)-296.4(1.)]TJ /F6 1 Tf 4.077 0 TD [(Schematic)-297.1(illustration)-290.3(of)-295.7(constructing)-296.7(complex)-292.8(organoids)-292.2(with)-293.1(adult)-294.4(stem)-294.2(cells)-295.4(and)-292.1(pluripotent)-294.9(stem)-294.2(cells,)-293(taking)]TJ -4.077 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[(system)-244.9(organoids.)]TJ 7.3823 0 0 6.5614 384.1511 521.2345 Tm (60)Tj 9.843 0 0 9.843 393.9023 516.8691 Tm [(Stem)-242.9(cells)-244(are)-241.7(specialized)-248.5(self-renewing)]TJ -8.2133 -1.1116 TD [(and)-511.3(multipotent)-504.7(cell)-508.1(types)-507.7(that)-506.9(contribute)-509(to)-503.4(turnover)-505.4(of)]TJ T* [(tissue)-289.9(ontogeny,)-284.2(growth,)-287.5(and)-280.9(lifelong)-286.8(regeneration.)-292.2(Figure)-284.2(3)]TJ 0 -1.1174 TD [(shows)-449.5(the)-448.1(currently)-450.2(widely)-446.6(recognized)-453.5(murine)-446.8(osteogenic)]TJ 0 -1.1116 TD [(stem)-294.7(cells,)-287.8(including)-287.9(BMSCs)-290.4(and)-292.4(SSCs,)-284.8(and)-292.4(illustrates)-291.1(their)]TJ T* [(main)-335.2(distribution)-336.2(areas)-337.1(within)-336.2(the)-332.9(bone)-333.9(\(Fig.)-336.1(3\).)]TJ /F3 1 Tf 8.9663 0 0 8.9663 323.0361 440.107 Tm (BMSCs.)Tj /F6 1 Tf 9.843 0 0 9.843 365.896 440.107 Tm [(BMSCs)-457.4(are)-460.6(nonhematopoietic)-467.5(ASCs)-461(in)-457.3(bone)]TJ -5.368 -1.1116 TD [(marro)-7.8(w)-445.5(and)-448(can)-449.1(differentiate)-451.4(into)-448.2(various)-446.5(tissues)-446.3(such)-448.9(as)]TJ T* [(bone,)-337.3(fat,)-336.3(and)-338.5(cartilage)]TJ /F7 1 Tf 9.7685 0 TD [(in)-336.4(vivo)]TJ /F6 1 Tf 3.1333 0 TD (and)Tj /F7 1 Tf 1.7798 0 TD [(in)-336.4(vitro)]TJ /F6 1 Tf 3.0123 0 TD [(.)-337.5(I)0(t)-336.4(i)0(s)-336.2(conrmed)]TJ -17.6938 -1.1116 TD [(that)-449.3(several)-448.8(murine)-446.8(BMSC)-445(subtyp)-6.5(es)-446.7(have)-444.5(the)-448.1(capacity)-445.7(to)]TJ 0 -1.1174 TD [(self-renew)-591.2(during)]TJ /F7 1 Tf 8.0118 0 TD [(in)-584.1(vivo)]TJ /F6 1 Tf 3.6229 0 TD [(sequential)-590.7(transplantation)-590.6(and)]TJ -11.6346 -1.1116 TD [(undergo)-377.5(trilineage)-377.9(differentiation.)]TJ 7.3823 0 0 6.5614 449.2345 389.707 Tm (6164)Tj 9.843 0 0 9.843 471.4014 385.3416 Tm [(These)-377.5(cell)-375.6(types)-375.2(in-)]TJ -16.0869 -1.1116 TD [(clude)-664.6(P)]TJ /F5 1 Tf 3.3925 0 TD (a)Tj /F6 1 Tf .5529 0 TD 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[(cell)-698.2(populations)-693.1(that)-691.2(had)-689.9(transcriptomic)-694.3(similaritie)-10.5(s)-686.3(t)0(o)]TJ T* (LepR)Tj /F11 1 Tf 7.3823 0 0 6.5614 335.055 126.7086 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 341.9716 122.3433 Tm [(cells,)-293.6(osteoblasts,)-299.9(chondrocytes,)-296.5(and)-292.4(broblasts)-294.1(were)]TJ -2.9374 -1.1174 TD [(placed)-361.6(downstream.)]TJ 7.3823 0 0 6.5614 393.0518 115.7102 Tm (67)Tj 9.843 0 0 9.843 403.9936 111.3448 Tm [(However,)-355(it)-359.6(is)-353.5(inappropriate)-358(to)-359.4(draw)]TJ -9.2386 -1.1116 TD [(any)-384.6(conclusion)-376.3(based)-382(only)-378.9(on)-382.3(the)-379(results)-377.3(of)-382.3(single-cell)-378.8(se-)]TJ T* [(quenc)-7.8(ing,)-283.3(and)-286.7(the)-286.8(data)-289.1(need)-289(to)-284.5(be)-291.3(further)-285.4(veried)-292.2(by)-284.4(further)]TJ 0 -1.1174 TD [(resear)-9.8(ch.)]TJ 1.0137 -1.1116 TD [(With)-577.1(regard)-579.1(to)-578.3(human)-572.4(BMSCs,)-581.8(a)-572.5(subendothelial)-583.7(cell)]TJ -1.0137 -1.1116 TD [(population)-519.4(expressing)-514.1(melanoma-associated)-521.1(cell)-513.8(adhesion)]TJ /F9 1 Tf 8.9663 0 0 8.9663 62.022 749.6502 Tm (548)Tj 48.541 0 TD [(QING)-334.9(ET)-338.7(AL.)]TJ ET endstream endobj 68 0 obj <>stream 0 0 0 rg /GS1 gs 119.282 56.58 .22676 697.323 re f 158.003 56.58 .22678 697.323 re f BT /F10 1 Tf 0 9.845 -9.845 -0 113.8393 245.3669 Tm 0 Tc 0 Tw (Table)Tj /F6 1 Tf 3.0002 0 TD (1.)Tj /F10 1 Tf 1.2496 0 TD [(Skeletal)-335.6(System)-334.6(Organoids)-332.7(and)-337.8(Their)-329.9(Fabrication)-334.2(Protocols)]TJ /F7 1 Tf -23.4258 -2.8562 TD (Fabrication)Tj 0 -1.0135 TD [(schemes)-6189.8(Noteworthy)-336.9(details)-5139.3(Materials)-5435.8(Cell)-334.7(source)]TJ 40.9087 2.027 TD [(Adherent)-333.9(culture)]TJ -1.0769 -1.0135 TD [(carried)-337.3(out)-331.4(before)-336.5(3D)]TJ 3.0808 -1.0135 TD (culture)Tj 8.4248 1.0135 TD (Long-term)Tj .0806 -1.0135 TD [(expansion)-3023(Culture)-337.5(products)-1414(Reference)]TJ /F6 1 Tf -51.418 -1.9061 TD (Suspension)Tj .9905 -1.0135 TD (culture)Tj 6.2998 1.0135 TD [(Cells)-335.5(were)-335.4(centrifuged)-337.2(to)]TJ .9905 -1.0135 TD [(generate)-335.3(pellets)]TJ 11.7589 1.0135 TD [(No)-7897.8(Human)-330.6(BMSCs)-335.6(from)]TJ 10.1062 -1.0135 TD [(iliac)-336.1(crest)-335.5(marrow)]TJ 0 -1.0077 TD (aspirates)Tj 9.3346 2.0212 TD [(2025)-333.2(days)-333.2(in)-330.4(adherent)]TJ .9905 -1.0135 TD (culture)Tj 9.3346 1.0135 TD [(Not)-332.2(mentioned)-1826.7(Chondro-osseous)]TJ 8.8854 -1.0135 TD (organoids)Tj 9.8817 1.0135 TD (46)Tj -61.2824 -3.0348 TD [(Cells)-335.5(were)-335.4(seeded)-337.5(in)-330.4(round-)]TJ .9905 -1.0135 TD [(bottom)-335.7(microwells)]TJ 11.7589 1.0135 TD [(PEG)-338.8(hydrogel)]TJ .9962 -1.0135 TD (microwells)Tj 8.1195 1.0135 TD [(Purchased)-334.6(human)]TJ .9905 -1.0135 TD [(MSCs)-333.7(and)]TJ 0 -1.0077 TD [(endothelial)-337.6(cells)]TJ 9.3346 2.0212 TD [(No)-9107.1(Not)-332.2(mentioned)-1826.7(Human)-250(bone)-241.2(marrow)]TJ 19.2105 -1.0135 TD (organoids)Tj 9.8817 1.0135 TD (50)Tj 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1.0135 TD (53)Tj -61.2824 -2.0212 TD [(Cartilage)-336.4(nodules)-336.3(were)]TJ .9905 -1.0135 TD [(formed)-336.5(by)-330.2(cell)]TJ T* [(differentiation)-336(in)-336.2(adherent)]TJ T* (culture)Tj 11.7589 3.0405 TD [(No)-7897.8(Human)-330.6(iPSCs)-4612.4(Cells)-335.5(differentiate)-335(for)]TJ 20.4313 -1.0135 TD [(14)-330.2(days)]TJ 9.3346 1.0135 TD [(Not)-332.2(mentioned)-1826.7(Cartilaginous)]TJ 8.8854 -1.0135 TD (organoids)Tj 9.8817 1.0135 TD (54)Tj -68.5727 -4.0483 TD [(Using)-334(hydrogel)]TJ .9905 -1.0135 TD (scaffold)Tj 6.2998 1.0135 TD [(Hydrogels)-293.1(containing)-289.2(BMSCs)]TJ .9905 -1.0135 TD [(were)-335.4(implanted)]TJ T* [(subcutaneously)-334.1(to)-336.2(form)]TJ 0 -1.0077 TD (organoid)Tj 11.7589 3.0348 TD [(Hybrid)-335.2(TG-PEG/HA)]TJ .9962 -1.0135 TD (hydrogels)Tj 8.1195 1.0135 TD [(Unsorted)-332.4(human)]TJ .9905 -1.0135 TD [(BMSCs)-335.6(from)-333.4(bone)]TJ T* [(marrow)-337.3(aspirates)]TJ 9.3346 2.027 TD [(Not)-332.2(mentioned)-4256.8(Not)-332.2(mentioned)-1826.7(Bone)-334.4(marrow)]TJ 19.2105 -1.0135 TD (organoid)Tj 9.3807 1.0135 TD (103)Tj -60.7814 -4.0483 TD [(Cells)-335.5(were)-335.4(seeded)-337.5(on)-330.2(porous)]TJ .9905 -1.0135 TD [(3D)-331(silk)-338.1(broin)-334.2(scaffolds)]TJ T* [(and)-332.4(cultured)-338.8(within)-329.8(a)]TJ 0 -1.0077 TD [(spinning)-330.6(bioreactor)]TJ 11.7589 3.0348 TD [(Silk)-303.6(broin)-305.4(scaffolds)-615.7(Unsorted)-332.4(human)]TJ 10.1062 -1.0135 TD [(BMSCs)-249.3(isolated)-246.7(from)]TJ T* [(bone)-333.4(marrow)]TJ 9.3346 2.027 TD [(Cells)-335.5(were)-335.4(cultured)-333.1(to)]TJ .9905 -1.0135 TD [(passage)-338.3(4)]TJ 9.3346 1.0135 TD [(Not)-332.2(mentioned)-1826.7(Woven)-336.5(bone)]TJ 8.8854 -1.0135 TD (organoids)Tj 9.8817 1.0135 TD (49)Tj -61.2824 -4.0483 TD [(Cells)-335.5(were)-335.4(resuspended)-336.5(in)]TJ .9905 -1.0135 TD (Matrigel)Tj 11.7589 1.0135 TD [(Matrigel)-5660.7(Mixed)-335.3(MSCs)-333.7(and)]TJ 10.1062 -1.0135 TD 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ثyUWتӛvlUnlU6*OozrQT^ٲ$ޛBB1ZlS'rt桾\H;hGتӛk6*0E4?LwWZ> z|'fޛ`2텩Mq*ԦP?FI]Q_` _96'omAYX2M MrzXdsqd$u~_k)=[Kh$ F!OFx$Ll \3ʛU6*a/V&.v[3U7qN vJSl$\F ;BQ.C.(x#l0+^W6GfZ #fdKȶ, 4NT߂D$b.8@#hD(/%Y cf&@*iL$\D,?CIvӶ;UH_O ت6*o+v m"Z4nǓGd;uTnMaO(]k`H H2[K/E9=C$z#9T7+h=E&^FHcA}J9 Zĺ0y @!Xe0rʤU轻WWЭ|eg=Ԓ]3.(UyNV;z+ak-ܱJ 3>ث*厹iZީsmy :vl^2^X\YŸ1Ti_]^K]5!_GxG׭N-Fh7 V 1yWfh0DIT. wf=*zE{H'9Z&ٔ}s,xdGqmI%V/3_jZx<%Cf'P4t5z|q7g9WQ|jUcl$:W3 8$ٶ@K x^w򮹪hy;V6ߣt3_z2H'6Zl v\}!#Oi j^3G0HpLت ߧ$YTUhї%XUiPG|ث͊;?ot4uKWDDQgmXWyKOuzuo͛23S}bh$^Rk [fOmi]{W#'F[mso|$3\OrjjlUҴI;$dZA\ث8_ZZaS3+AkԪԑe`۰<0h-{r 8bv>ӮlU s[~a?H͜Y5pknYJACf]vlUٱWf]vlUٱWf]vlUٱWf]vlUٱWf]vlUٱWf]v endstream endobj 83 0 obj <>stream /GS1 gs BT /F6 1 Tf 9.843 0 0 9.843 62.022 725.1588 Tm 0 0 0 rg 0 Tc 0 Tw [(molecule)-503.4(\()-81.7(MCAM)-8.1(/CD146\))-499.3(has)-499.6(been)-502.1(identied)-502(in)-503.4(human)]TJ 0 -1.1174 TD [(bone)-351.2(marrow)-355.4(stroma.)]TJ 7.3823 0 0 6.5614 148.0818 718.5825 Tm (73)Tj 9.843 0 0 9.843 158.9669 714.1605 Tm [(The)-351.2(generation)-354.6(of)-347.7(heterotopic)-355.8(ossi-)]TJ -9.8491 -1.1116 TD [(cles)-392.6(within)-388(bone)-391.5(marrow)-395.7(and)-384.6(establishme)-11.1(nt)-388.2(of)-388.1(ectopic)-391.6(he-)]TJ T* [(matopoietic)-501.2(microenvironment)-502(were)-497.1(demonstrated)-501.8(in)-491.9(that)]TJ T* [(study.)-360.2(CD271)-358.8(has)-361.4(been)-363.8(identied)-358(to)-359.4(be)-360.5(a)-359.4(marker)-361.4(of)-359.3(a)-359.4(sub-)]TJ 0 -1.1174 TD [(population)-329.4(of)-319(self-renewing)-329.7(and)-327(PSCs)-327.5(in)-319.1(bone)-328.1(marrow.)]TJ 7.3823 0 0 6.5614 284.485 674.7021 Tm (74,75)Tj 9.843 0 0 9.843 62.022 659.3951 Tm [(One)-304.8(subpopulation)-303.7(that)-299.6(was)-304.6(specically)-304.8(labeled)-305.2(by)-301.7(CD271)]TJ /F11 1 Tf 7.3823 0 0 6.5614 297.0708 663.7605 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 62.022 648.4534 Tm (Thy1)Tj /F11 1 Tf 7.3823 0 0 6.5614 82.8283 652.8187 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 89.4614 648.4534 Tm (VCAM-1)Tj 7.3823 0 0 6.5614 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[(se)14.1(lf)20.7(-r)15.1(en)19.7(ew)13.1(in)20.7(g)-214.2(p)0(o)15.1(s)0(t)20.9(n)0(a)13.9(t)0(a)19.6(l)-214.3(mS)14(SC)20(s.)]TJ 7.3823 0 0 6.5614 185.2157 540.4534 Tm (79)Tj 9.843 0 0 9.843 194.6267 536.088 Tm [(Fu)15.2(rt)15(he)19.7(r)-214.2(e)0(x)13.9(p)0(l)20.7(o)0(r)15.1(a)0(t)19.5(i)0(o)15(n)-214.2(of)]TJ /F7 1 Tf 8.8469 0 TD [(bo)15.1(na)]TJ -22.3189 -1.1174 TD [(d)15.1(e)]TJ /F6 1 Tf 1.9525 0 TD [(hu)20.8(ma)12.7(n)-519.5(s)0(k)15.3(e)0(l)19.5(e)0(t)13.8(a)0(l)-510.3(s)0(t)15.1(e)0(m)-511.4(c)0(e)12.8(l)0(l)20.6(s)-525(\(h)20.9(SS)15.4(Cs)14.2(\))-519.4(r)0(e)14(v)0(e)19.7(a)0(l)19.5(e)0(d)-516(t)0(h)15(a)0(t)-510.3(t)0(h)15(e)]TJ -1.9525 -1.1116 TD [(PD)14.4(PN)]TJ /F11 1 Tf 7.3823 0 0 6.5614 86.6834 518.5132 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 93.0897 514.1479 Tm [(CD)13.2(14)20.8(6)]TJ /F11 1 Tf 7.3823 0 0 6.5614 121.096 518.5132 Tm (-)Tj /F6 1 Tf 9.843 0 0 9.843 127.5023 514.1479 Tm [(CD)18.9(73)]TJ /F11 1 Tf 7.3823 0 0 6.5614 150.633 518.5132 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 157.0393 514.1479 Tm [(CD)13.2(16)20.8(4)]TJ /F11 1 Tf 7.3823 0 0 6.5614 184.9889 518.5132 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 191.4519 514.1479 Tm [(ce)18.5(ll)-232.8(su)21(bp)15.1(op)20.8(ul)14.9(at)19.5(io)15(n)-237.2(e)0(x)19.7(h)0(i)14.9(b)0(i)20.7(t)0(e)13.8(d)]TJ -13.1494 -1.1116 TD [(th)15(e)-249.9(g)0(r)20.9(e)0(a)12.8(t)0(e)19.6(s)0(t)-244(h)0(o)20.8(m)0(o)19.6(l)0(o)15(g)0(y)-238.3(i)0(n)-244.2(t)0(e)19.6(r)0(m)13.9(s)-248.6(of)-238.3(ge)13.9(ne)-239.5(ex)13.9(pr)20.9(es)14.1(si)20.9(on)-244.1(to)-238.5(mS)19.8(SC)14.2(s,)]TJ 0 -1.1174 TD [(an)13.9(d)-392.8(t)0(h)15(e)0(i)19.5(r)-398.5(se)19.9(lf)15(-r)20.9(en)13.9(ew)18.8(al)-389.4(ca)18.5(pa)13.9(ci)19.5(ty)-388.2(an)13.9(d)-392.8(p)0(l)14.9(u)0(r)20.9(i)0(p)15(o)0(t)20.7(e)0(n)13.9(c)0(y)-383.5(w)0(e)13(r)0(e)-389.2(v)0(a)19.7(l)0(i)14.8(-)]TJ 0 -1.1116 TD [(da)13.9(te)19.6(d)-657.7(t)0(h)15(r)0(o)20.9(u)0(g)15.1(h)-657.7(el)19.5(eg)13.9(an)19.7(t)-657.8(c)0(o)13.9(n)0(t)20.7(i)0(n)15(u)0(o)20.8(u)0(s)-652.9(t)0(r)20.7(a)0(n)13.9(s)0(p)21(l)0(a)13.8(n)0(t)20.7(a)0(t)13.8(i)0(o)20.7(n)-657.7(ex)13.9(pe)19.7(ri)15(-)]TJ T* [(me)18.4(nt)14.9(s.)]TJ 7.3823 0 0 6.5614 87.5338 474.6896 Tm (80)Tj 9.843 0 0 9.843 98.4189 470.3243 Tm [(Bo)19.8(th)-353.7(mS)19.8(SC)14.2(s)-358(a)0(n)13.9(d)-358.2(hS)15.2(SC)20(s)-363.8(a)0(r)19.7(e)-365.1(lo)20.7(ca)12.8(te)19.6(d)-364(i)0(n)-347.9(t)0(h)15(e)-359.4(gr)15.1(ow)20(th)]TJ -3.6977 -1.1174 TD [(pl)15(at)19.5(e)-278.7(a)0(n)13.9(d)-277.6(ca)18.5(n)-277.6(g)0(e)13.9(n)0(e)19.7(r)0(a)14(t)0(e)-268.4(p)0(r)20.9(o)0(g)15.1(e)0(n)19.7(i)0(t)14.8(o)0(r)-267.1(c)0(e)12.8(l)0(l)20.6(s)-277.4(of)-272.9(bo)20.8(ne)13.9(,)-274.1(c)0(a)12.8(r)0(t)20.7(i)0(l)14.8(a)0(g)19.7(e)-278.7(an)13.9(d)]TJ 0 -1.1116 TD [(st)15.1(ro)20.9(ma)12.7(,)-285.7(b)0(u)20.8(t)-289.2(no)15.1(t)-289.2(f)0(a)19.7(t)0(.)-281.1(I)0(n)-278.6(a)0(d)13.9(d)0(i)20.7(t)0(i)14.8(o)0(n)20.8(,)-291.4(re)19.7(st)15.1(in)20.7(g)-289.1(c)0(h)19.7(o)0(n)15.1(d)0(r)15.1(o)0(c)19.7(y)0(t)20.7(e)0(s)-285.4(t)0(h)20.7(a)0(t)-280(e)0(x)13.9(-)]TJ T* [(pr)15.1(es)19.9(s)-219.8(p)0(a)19.7(r)0(a)14(t)0(h)20.7(y)0(r)15.1(o)0(i)14.9(d)-214.2(ho)15.1(rm)19.7(on)15.1(e-)19.7(re)14(la)19.6(te)13.8(d)-214.2(p)0(r)15.1(o)0(t)20.7(e)0(i)13.8(n)-220(we)18.8(re)-210.7(id)15(en)19.7(ti)14.8(e)19.8(d)-220(a)0(s)-210.5(a)]TJ T* [(su)15.3(bg)20.8(ro)15.1(up)-261.4(of)-261.4(SS)15.4(Cs)-262.2(an)19.7(d)-271.8(p)0(r)15.1(o)0(g)20.8(e)0(n)19.7(i)0(t)14.8(o)0(r)-261.4(c)0(e)12.8(l)0(l)20.6(s)-271.6(in)-261.5(th)15(e)-267.2(g)0(r)15.1(o)0(w)20(t)0(h)-267.3(p)0(l)20.7(a)0(t)13.8(e)0(.)]TJ 7.3823 0 0 6.5614 291.4582 430.866 Tm -.0162 Tc (81)Tj 9.843 0 0 9.843 72 415.5022 Tm 0 Tc [(Another)-332.4(group)-328(of)-330.5(SSCs)-333.2(were)-335.9(identied)-329.3(around)-336(the)-327.1(blood)]TJ -1.0137 -1.1116 TD [(vessels.)-278(These)-279.5(were)-278.3(CD45)]TJ /F11 1 Tf 7.3823 0 0 6.5614 166.7338 408.9259 Tm (-)Tj /F6 1 Tf 9.843 0 0 9.843 173.537 404.5605 Tm (CD31)Tj /F11 1 Tf 7.3823 0 0 6.5614 197.0078 408.9259 Tm (-)Tj /F6 1 Tf 9.843 0 0 9.843 203.811 404.5605 Tm (PDGFR)Tj /F5 1 Tf 3.2082 0 TD (a)Tj /F11 1 Tf 7.3823 0 0 6.5614 240.8314 408.9259 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 247.5779 404.5605 Tm (Sca1)Tj /F11 1 Tf 7.3823 0 0 6.5614 266.7968 408.9259 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 273.5432 404.5605 Tm (CD24)Tj /F11 1 Tf 7.3823 0 0 6.5614 297.0708 408.9259 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 62.022 393.6188 Tm [(cells,)-345.4(which)-347.5(were)-341.6(termed)-344.3(as)-343(perivascular)-351.9(SSCs.)-342.4(In)-342(addition)]TJ 0 -1.1174 TD [(to)-468.9(differentiating)-469.8(into)-465.4(bone)-466.3(and)-471(cartilage,)-467.6(these)-468.5(cells)-468.7(had)]TJ 0 -1.1116 TD [(adipogenic)-446.7(potential)-450.5(and)-442.2(formed)-446.7(bone)-443.3(marrow)-447.5(adipocytes)]TJ T* [(after)-324.7(transplantation)]TJ /F7 1 Tf 8.4495 0 TD [(in)-319.1(vivo)]TJ /F6 1 Tf 2.7762 0 TD (.)Tj 7.3823 0 0 6.5614 175.011 365.1023 Tm (82)Tj 9.843 0 0 9.843 185.5558 360.7369 Tm [(These)-319.9(formed)-320(a)-319(distinct)-321.3(SSC)]TJ -12.5504 -1.1174 TD [(subpopulation)-286.4(from)-287.8(the)-281.1(primarily)-289.1(identied)-288.9(mSSCs,)-281.4(and)-286.7(did)]TJ 0 -1.1116 TD [(not)-331.7(overlap)-338.4(with)-333.8(each)-336.2(other.)]TJ 7.3823 0 0 6.5614 175.8614 343.1621 Tm (83)Tj 9.843 0 0 9.843 72 327.855 Tm [(The)-420.4(distribution)-422.6(of)-422.6(SSCs)-419.6(was)-419.8(also)-426(found)-420.2(in)-417(periosteum)]TJ -1.0137 -1.1174 TD [(and)-390.4(fracture)-398.2(calluses.)]TJ 7.3823 0 0 6.5614 148.4787 321.2787 Tm (8486)Tj 9.843 0 0 9.843 170.8157 316.8566 Tm [(Debnath)-391.2(et)-395.1(al.)-387.1(identied)-398.4(perios-)]TJ -11.0529 -1.1116 TD [(teal)-467.8(stem)-467.5(cells)-468.7(that)-466.6(are)-466.4(present)-470.7(in)-463.1(murine)-469.9(long)-465.3(bone)-466.3(and)]TJ T* (skull.)Tj 7.3823 0 0 6.5614 83.6787 299.3385 Tm (84)Tj 9.843 0 0 9.843 95.5842 294.9731 Tm [(These)-463.9(were)-462.6(specically)-466.1(labeled)-466.5(with)-460.6(cathepsin)-468.5(K)]TJ -3.4097 -1.1116 TD [(\(Ctsk\))-399.2(and)-396.1(located)-397.4(at)-400.9(the)-396.3(apex)-398.4(of)-393.8(the)-396.3(differentiation)-400.7(level.)]TJ 0 -1.1174 TD [(Rigorous)-418.7(continuous)-415.4(transplantation)-423.6(experiments)-420.1(validated)]TJ 0 -1.1116 TD [(the)-332.9(pluripotency)-338.5(and)-332.8(self-renewal)-341.3(capacity)-336.3(of)-336.2(these)-336.1(cells.)]TJ 1.0137 -1.1116 TD [(One)-304.8(other)-306.3(periosteum)-308.3(SSC)-303.5(subpopulation)-309.4(with)-305(high)-304(bone)]TJ -1.0137 -1.1174 TD [(regenerative)-514.6(potential)-513.9(was)-506.2(chara)-9(cterized)-509(by)-509.1(Prx1)-511.1(expres-)]TJ 0 -1.1116 TD (sion.)Tj 7.3823 0 0 6.5614 80.9008 233.5747 Tm (85)Tj 9.843 0 0 9.843 91.1055 229.2094 Tm [(These)-291.1(could)-289(mature)-286.7(into)-286.9(osteoblasts,)-294.2(adipocytes,)-288.7(and)]TJ -2.9547 -1.1116 TD [(chondrocyte)-344(lineages)]TJ /F7 1 Tf 8.9275 0 TD [(in)-336.4(vitro)]TJ /F6 1 Tf 3.0181 0 TD [(,)-337.5(and)-338.5(possessed)-339.8(higher)]TJ /F7 1 Tf 9.5496 0 TD [(in)-336.4(vivo)]TJ /F6 1 Tf -21.4952 -1.1174 TD [(regenerative)-243.9(potential)-237.4(than)-236.1(BMSCs.)-236.2(The)-236(combination)-240.7(of)-232.6(MX)]TJ 0 -1.1116 TD [(dynamin)-419.2(like)-414.8(GTPase)-425.3(1)-415.8(\()-81.7(MX1\))-420.9(and)-413.4(alpha)-422.7(smooth)-416.7(muscle)]TJ T* [(Actin)-399.5(\()]TJ /F5 1 Tf 2.9605 0 TD (a)Tj /F6 1 Tf .5529 0 TD [(SMA\))-397.7(expression)-404.7(can)-397.3(be)-400.8(used)-402.8(to)-394(selectively)-408.8(label)]TJ -3.5134 -1.1116 TD [(SSCs.)-330.9(MX1)]TJ /F11 1 Tf 7.3823 0 0 6.5614 109.7007 178.8094 Tm (+)Tj /F5 1 Tf 9.843 0 0 9.843 117.0141 174.444 Tm (a)Tj /F6 1 Tf .5529 0 TD (SMA)Tj /F11 1 Tf 7.3823 0 0 6.5614 143.6598 178.8094 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 150.9732 174.444 Tm [(cells)-336.2(are)-333.9(present)-332.5(in)-330.6(the)-327.1(periosteum)-337.1(of)]TJ -9.037 -1.1174 TD [(adult)-369.8(mice,)-368.6(and)-361.6(had)-367.3(the)-367.5(capacity)-370.8(for)-366.1(colony)-365(formation)-369.6(and)]TJ 0 -1.1116 TD [(trilineage)-423.9(differentiation)-423.7(potential)]TJ /F7 1 Tf 14.3359 0 TD [(in)-417(vitro)]TJ /F6 1 Tf 3.093 0 TD [(.)-418.1(These)-423.5(cells)-422.6(re-)]TJ -17.4289 -1.1116 TD [(tained)-423.9(the)-419.3(expression)-427.8(of)-422.6(CD140a)]TJ /F11 1 Tf 7.3823 0 0 6.5614 198.0283 145.9275 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 206.3055 141.5621 Tm [(and)-419.2(CD105)]TJ /F11 1 Tf 7.3823 0 0 6.5614 253.0771 145.9275 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 261.2976 141.5621 Tm [(after)-422.6(con-)]TJ -20.2454 -1.1174 TD [(tinuous)-336.1(transplantation)-337.2(into)-333(skull)-333.9(defect)-338.5(sites.)]TJ 7.3823 0 0 6.5614 244.97 134.9291 Tm (86)Tj /F12 1 Tf 8.9663 0 0 8.9663 62.022 105.9023 Tm [(Endogenous)-336.3(and)-329.4(exogenous)-333.5(signals)]TJ /F6 1 Tf 9.843 0 0 9.843 72 89.4614 Tm [(Successful)-267.5(organoid)-263.5(culture)-269.4(lies)-264.8(in)-267.3(knowledge)-266.7(of)-261.4(the)-263.8(stem)]TJ -1.0137 -1.1174 TD [(cell)-606(microenvironment)-611.4(and)-609.2(their)-605.9(lineage)-610.5(fate)-605.8(regulation)]TJ 0 -1.1116 TD [(mechanisms.)-584.6(The)-587.4(derivation)-583.9(of)-578.1(organoids)-587(depends)-580.1(part-)]TJ T* [(ly/entirely)-533.4(on)-526.3(endogenous)-531.5(and)-528.6(exogenous)-530.4(signals,)-530.5(which)]TJ 25.504 67.9242 TD [(mimic)-331.9(the)-327.1(niche)-330.5(factors)-332.5(encountered)-333.7(by)-330.5(stem)-329.3(cells.)-328.1(Current)]TJ 0 -1.1174 TD [(studie)-7.8(s)-242.8(have)-254.4(provided)-246.2(some)-248.5(cues)-248.4(to)-250(the)-246.5(factors)-251.8(that)-247.7(inuence)]TJ 0 -1.1116 TD [(stem)-340.8(cell)-335.3(lineage)-334(fate.)]TJ 1.0137 -1.1116 TD [(Granulocyte)-500.5(colony-stim)-8.9(ulating)-498.8(factor)-498.6(inhibits)-498.7(the)-499.9(pro-)]TJ -1.0137 -1.1116 TD [(liferation)-393.9(of)-388.1(Nestin)]TJ /F11 1 Tf 7.3823 0 0 6.5614 390.9542 685.7006 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 398.8912 681.3352 Tm [(MSCs)-391.9(and)-390.4(downregulates)-393.1(osteogenic-)]TJ -8.7202 -1.1174 TD [(specic)-650.3(differentiation)-648.4(genes)-641.2(in)-647.4(cells.)-644.9(By)-642.6(contrast,)-648.2(the)]TJ /F7 1 Tf 0 -1.1116 TD [(in)-474.6(vivo)]TJ /F6 1 Tf 3.404 0 TD [(administration)-474.4(of)-468.7(parathyroid)-475.4(hormone)-470.9(stimulated)]TJ -3.404 -1.1116 TD [(bone)-345.4(marrow)-338.1(Nestin)]TJ /F11 1 Tf 7.3823 0 0 6.5614 395.3196 652.8187 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 402.7463 648.4534 Tm [(MSCs)-340.1(to)-342.2(proliferate)-344.2(and)-338.5(differentiate)]TJ -9.1119 -1.1174 TD [(into)-453.9(the)-459.6(osteogenic)-458.1(lineage.)]TJ 7.3823 0 0 6.5614 427.2944 641.877 Tm (62)Tj 9.843 0 0 9.843 439.1432 637.455 Tm [(Phosphatase)-458.7(and)-453.7(tensin)-451.3(ho-)]TJ -12.8096 -1.1116 TD [(molog)-387(regulate)-392.7(the)-384.7(quiescence,)-388.8(maintenance,)-391.5(and)-384.6(differen-)]TJ T* [(tiation)-838.8(of)-837.3(LepR)]TJ /F11 1 Tf 7.3823 0 0 6.5614 385.0015 619.9368 Tm (+)Tj /F6 1 Tf 9.843 0 0 9.843 397.3605 615.5715 Tm [(cells.)-835(Specically,)-843.9(it)-837.6(can)-835(promote)]TJ -8.5647 -1.1174 TD [(differentiation)-452.5(into)-442.4(osteoblasts)-446.2(and)-442.2(inhibit)-446(adipogenic)-446.7(dif-)]TJ 0 -1.1116 TD [(ferentiatio)-10.3(n.)]TJ 7.3823 0 0 6.5614 361.1905 597.9967 Tm (64)Tj 9.843 0 0 9.843 372.0188 593.6313 Tm [(Jun-B)-346.1(is)-347.7(a)-347.8(very)-351.1(important)-346.7(factor)-348.8(required)-351(by)]TJ -5.9901 -1.1116 TD [(stromal)-361.5(cells)-365(to)-353.7(proliferate)-367.2(and)-355.8(differentiate)-365(into)-356(the)-361.7(osteo-)]TJ T* [(genic)-347.8(lineage.)-349(It)-347.9(controls)-344.1(BMSC)-347(fate)-346.6(during)-346.5(bone)-345.4(develop-)]TJ 0 -1.1174 TD [(ment.)-321.4(Knockout)-322(of)-319(Jun-B)-323.1(had)-321.2(been)-317.8(demonstrated)-329(to)-319.1(lead)-317.9(to)]TJ 0 -1.1116 TD [(the)-511.4(decrease)-509.7(of)-503.3(osteoprogenitor)-514.3(cell)-508.1(pools,)-504.2(H-type)-509.8(blood)]TJ T* [(vessels,)-318.3(and)-309.7(arteries)-317.6(in)-307.6(bone,)-314.3(while)-313.1(increasing)-314(adipocytes.)]TJ 7.3823 0 0 6.5614 544.8187 543.2313 Tm (72)Tj 9.843 0 0 9.843 323.0361 527.8676 Tm -.0157 Tc [(De)5.1(cre)5.1(a)-.7(se)5.1(d)-373.9(p).5(r)6.3(o).5(li)6.1(fer)6.3(a)-.7(ti)6.1(on)-373.9(a)5.1(n).5(d)-373.9(i)6.1(ncr)6.3(e)-.7(a)5.1(s).7(ed)-368.1(apo)6.2(p).5(to)6.2(tic)-369.3(act)6.1(ivi)6.1(t).4(y)-368.1(i).4(n)]TJ -1.0137 -1.1116 TD -.0165 Tc [(diabet)5.3(ic)-214.6(mi)5.3(ce)-214.6(SS)5.6(Cs)-213.2(wer)5.5(e)-214.6(rev)5.4(e)-1.5(rsed)-213.4(upo)5.4(n)-213.4(l)-.5(ocal)-213.6(deliv)5.4(ery)-213.4(o)-.3(f)-213.4(I)-.3(n)5.4(d)-.3(ian)]TJ T* -.0157 Tc [(hed)6.2(g).5(eh)6.2(og.)]TJ 7.3823 0 0 6.5614 352.2896 510.3495 Tm -.0239 Tc (87)Tj 9.843 0 0 9.843 363.2314 505.9841 Tm -.0157 Tc [(P)6.4(a)-.7(ra)5.1(cri)6.1(n).5(e)-392.3(f).5(ac)5.1(tor)6.3(s)-396.7(se)5.1(cr)6.3(ete)5.1(d)-396.9(b)6.2(y)-396.9(S)6.4(c)-.7(hw)5.4(ann)-391.2(c)5.1(ell)6.1(s)-396.7(a)5.1(r).5(e)]TJ -5.0973 -1.1174 TD -.0159 Tc [(also)-477.8(con)6(s).5(id)6(ere)4.9(d)-483.5(to)-483.5(h)6(a)-.9(ve)-478.9(pot)5.9(ent)5.9(i).2(al)-483.7(e)4.9(ffe)4.9(ct)5.9(s)-483.3(o).3(n)-483.5(m)4.8(SSC)5(s)-483.3(i).1(n)-483.5(t).2(h)6(e)]TJ 0 -1.1116 TD -.0153 Tc [(man)6.6(dib)6.6(l).7(e.)]TJ 7.3823 0 0 6.5614 350.7589 488.4093 Tm -.0239 Tc (88)Tj 9.843 0 0 9.843 361.7574 484.0439 Tm -.0156 Tc [(Va)5.2(scu)6.3(l).5(ar)-402.6(e)5.2(ndo)6.3(the)5.2(l).4(ia)5.2(l)-402.7(g).6(r)6.4(o).6(wt)6.2(h)-402.6(f).6(ac)5.2(tor)-402.6(\()6.4(VEG)5.5(F).7(\))-402.6(w)5.5(as)]TJ -4.9476 -1.1116 TD -.0165 Tc 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[(12.)-500.5(Sato)-415.8(T,)-416.9(Vries)-422.1(RG,)-415.4(Snippert)-418.1(HJ,)-415.8(et)-418.7(al.)-418.7(Single)-415.5(Lgr5)-419.5(stem)]TJ 1.7505 -1.1223 TD [(cells)-367(build)-365.5(crypt-villus)-367.3(structures)-365.3(in)-365.7(vitro)-361.7(without)-366.8(a)-364.2(mes-)]TJ 0 -1.1284 TD [(enchymal)-275.1(niche.)-270.9(Nature)-274.9(2009;459\(7244)14.7(\):262265;)-271(doi:)-273.9(10)]TJ 0 -1.1223 TD (.1038/nature07935)Tj -1.7505 -1.1284 TD [(13.)-500.5(Ashok)-782.1(A,)-782.5(Choudhury)-783(D,)-782.5(Fang)-782.3(Y,)-782.5(et)-784.6(al.)-784.7(Towards)]TJ 1.7505 -1.1223 TD [(manufacturing)-749.5(of)-746.7(human)-748.6(organoids.)-750.6(Biotechnol)-750.9(Adv)]TJ 0 -1.1284 TD [(2020;39:107460;)-335.9(doi:)-328.8(10.1016/j.biotechadv.2019.107460)]TJ -1.7505 -1.1223 TD [(14.)-500.5(Sc)23.1(hu)24.1(t)12.6(g)0(e)19.9(n)0(s)-347.1(F)15.3(,)-359.9(R)0(o)19.3(o)0(k)24.1(m)12.5(a)14(a)0(k)19.9(e)14(r)-362.3(M)12.8(B)13.4(,)-359.9(M)12.8(a)0(r)17.6(g)0(a)26(r)0(i)22.4(t)12.6(i)12.6(s)-359.1(T,)-349.8(et)-351.6(a)14(l)12.6(.)-359.9(T)0(u)22.2(b)0(u)24.1(-)]TJ 1.7505 -1.1284 TD [(l)12.6(o)0(i)24.7(d)0(s)-353.2(d)0(e)26(r)0(i)22.4(v)0(e)19.9(d)-360(f)15.8(r)0(o)21.8(m)-359.6(hu)24.1(ma)20.3(n)-360(a)0(d)19.9(u)0(l)24.7(t)-359.4(ki)24.7(dn)24.1(ey)-352.1(an)19.9(d)-360(u)0(r)27.9(i)0(n)18.6(e)-358.1(fo)21.8(r)-362.3(p)0(e)26(r)0(-)]TJ 0 -1.1223 TD [(s)12.9(o)0(n)24.1(a)0(l)20.5(i)12.6(ze)21.8(d)-536.9(d)0(i)24.7(s)12.9(ea)21.8(s)12.9(e)-535(mo)18.4(de)26(l)12.6(i)12.6(n)0(g)18(.)-530.7(Na)24.4(t)-536.3(B)13.4(io)18.6(t)12.6(e)14(ch)19.9(no)24.1(l)-536.3(2)0(0)24.1(1)0(9)24.1(;)12.6(37)24.1(\(3)21.8(\))15.8(:)]TJ 0 -1.1284 TD 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[(doi:)-334.9(10.1089/ten.tea.2020.0273)]TJ -1.7505 -1.1284 TD [(54.)-500.5(Tam)-274.5(WL,)-275(Men)14.4(des)-272.1(LF,)-273.5(Che)15(n)-286.8(X)0(K)14.9(,)-286.7(et)-278.4(al.)-272.4(Hum)16.7(an)-279(plu)12.3(rip)16.1(ot)12.5(ent)]TJ 1.7505 -1.1223 TD [(ste)15.1(m)-201(cel)16.1(l-d)16.1(eri)11.9(ved)-187.6(car)13.2(til)13.5(ag)13.8(ino)12.3(us)-194.6(org)15.5(an)13.8(oid)12.3(s)-194.4(pro)15.5(mot)12.8(e)-199.5(sca)16.4(ffo)13.2(ld)12.5(-)]TJ 0 -1.1284 TD [(fre)15.1(e)-193.4(hea)15.6(lin)12.9(g)-189.2(o)0(f)-185.6(cri)11.9(ti)13.1(cal)-185.1(siz)15.1(e)-193.4(l)0(o)12.5(n)0(g)-189.4(bon)17.9(e)-193.4(d)0(e)13.8(fec)13.2(ts.)-181.9(Ste)11.3(m)-188.8(Cel)15.5(l)-194.7(Res)]TJ 0 -1.1223 TD [(The)11.8(r)-270.8(2021)17.7(;1)12.5(2\(1)15.5(\):5)16.1(13;)-268.2(doi)18.4(:)-274(1)0(0)18(.118)17.8(6/)12.5(s13)12.7(287)17.9(-02)15.5(1-0)15.5(258)17.9(0-7)]TJ 25.263 70.8975 TD [(55.)-500.5(Li)-428.5(ML,)-428.5(Aggeler)-433.6(J,)-432.4(Farson)-430.2(DA,)-430.5(et)-430.9(al.)-430.9(Inuence)-433.8(of)-429.6(a)-431.3(re-)]TJ 1.7505 -1.1223 TD [(constituted)-464.3(basement)-475.4(membrane)-466.6(and)-468.2(its)-468(components)-465.4(on)]TJ 0 -1.1284 TD [(casein)-323.1(gene)-320.1(expression)-327(and)-321.8(secretion)-325.3(in)-316.9(mouse)-326.9(mammary)]TJ 0 -1.1223 TD [(epithelial)-438.1(cells.)-446.3(Proc)-436.9(Natl)-444.4(Acad)-443.6(Sci)-440(U)-440.9(S)-436.1(A)-440.9(1987;84\(1\):)]TJ 0 -1.1283 TD [(136140;)-330(doi:)-334.9(10.1073/pnas.84.1.136)]TJ -1.7505 -1.1223 TD [(56.)-500.5(Akkerman)-293.6(N,)-288.5(Deze)-298.4(LHK.)-292.2(Dawn)-294.6(of)-289.3(the)-290.7(organoid)-293.8(era:)-291.3(3D)]TJ 1.7505 -1.1284 TD [(tissue)-435.2(and)-431.6(organ)-440.3(cultures)-436.8(revolutionize)-435.4(the)-437.1(study)-432.3(of)-435.6(de-)]TJ 0 -1.1223 TD [(velopment,)-1283(disease,)-1280(and)-1279.4(regeneration.)-1290.8(Bioessays)]TJ 0 -1.1284 TD 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[(liver.)-408.5(C)-4.7(ell)-408(2015;)-5.5(160\(12\):299312;)-408(doi)-5.5(:)-401.9(1).1(0)-24.3(.1016/j)-23.8(.).1(cell)]TJ 0 -1.1223 TD 0 Tc (.2014.11.050)Tj -1.7505 -1.1284 TD [(59.)-500.5(Rossi)-295.9(G,)-294.6(Manfrin)-295.2(A,)-294.6(Lutolf)-296.7(MP.)-295.3(Progress)-297.8(and)-297.4(potential)-294.1(in)]TJ 1.7505 -1.1223 TD [(organoid)-367(research.)-375.6(Nat)-371.6(Rev)-375.4(Genet)-370(2018;19\(11\):671687;)]TJ 0 -1.1284 TD [(doi:)-328.8(10.1038/s41576-018-0051-9)]TJ -1.7505 -1.1223 TD [(60.)-500.5(Sipp)-234.7(D,)-239.7(Robey)-241.5(PG,)-236.6(Turner)-243.4(L.)-233.9(Clear)-241.3(up)-238.2(this)-242.5(stem-cell)-238.9(mess.)]TJ 1.7505 -1.1284 TD .0122 Tc [(Nature)-400.8(2018;5)-6.2(61\(77)-6.2(24\):)-5.7(455457;)-402.1(doi:)-402.1(1)-.1(0)-12.3(.1038/d415)-6.2(86-)]TJ 0 -1.1223 TD 0 Tc (018-06756-9)Tj -1.7505 -1.1284 TD [(61.)-500.5(Morikawa)-469.6(S,)-460.6(Mabuchi)-471.7(Y,)-465.3(Kubota)-469.6(Y,)-465.3(et)-467.5(al.)-461.4(Prospective)]TJ 1.7505 -1.1223 TD [(identication,)-490.4(isolation,)-496.4(and)-492.6(systemic)-494.2(transplantation)-495.2(of)]TJ 0 -1.1284 TD [(multipotent)-667.3(mesenchymal)-674.8(stem)-667.7(cells)-671.9(in)-664.5(murine)-671.2(bone)]TJ 0 -1.1223 TD .0183 Tc [(marrow.)-402.6(J)-407.9(Ex)6(p)-408.8(M).6(ed)-408.8(2)6(009;206\(11\):24832496;)-402.1(doi:)-402.1(1)-.1(0)]TJ 0 -1.1284 TD 0 Tc (.1084/jem.20091046)Tj -1.7505 -1.1223 TD [(62.)-500.5(Mendez-Ferrer)-473.5(S,)-472.7(Michurina)-479.8(TV,)-473.5(Ferraro)-470.6(F,)-472.8(et)-473.6(al.)-473.6(Me-)]TJ 1.7505 -1.1284 TD [(senchymal)-390.2(and)-388.9(haematopoietic)-398.5(stem)-387.1(cells)-391.3(form)-395.1(a)-388.6(unique)]TJ 0 -1.1223 TD [(bone)-261(mar)11.8(row)-260.5(nich)14.1(e.)-272.8(Na)12.2(ture)-262.7(2010)17.7(;46)12.3(6\(73)15.4(08\):)16(8298)17.6(34;)-262.1(doi)12.3(:)]TJ 0 -1.1284 TD (10.1038/nature09262)Tj -1.7505 -1.1223 TD [(63.)-500.5(Pinho)-308(S,)-314.2(Lacombe)-314(J,)-316.5(Hanoun)-311.6(M,)-310.5(et)-315(al.)-315.1(PDGFR)-313.3(alpha)-313.5(and)]TJ 1.7505 -1.1284 TD [(CD51)-385.1(mark)-390.9(human)-388.8(Nestin\()]TJ /F11 1 Tf 11.2897 0 TD (+)Tj /F6 1 Tf .5489 0 TD [(\))-392.8(sphere-forming)-389.9(mesenchy-)]TJ -11.8386 -1.1223 TD [(mal)-424.5(stem)-429.8(cells)-428(capable)-432(of)-423.5(hematopoietic)-430.7(progenitor)-430(cell)]TJ 9.2951 0 0 9.2951 331.7668 359.0361 Tm .0065 Tc [(e)6.5(x)-10.2(pans)-5(i)-5.3(on.)-396.1(J)-401.4(E)-1.6(xp)-402.3(M)6.5(e)-9.5(d)-396.2(2)-5.8(013)-5.8(;)6.5(2)-11.5(10\(7)-5.8(\))-2.1(:)6.5(1)-11.5(351)-5.8(13)-5.8(67)-5.8(;)-395.6(d).3(o)-5.8(i)6.5(:)-401.3(1)-5.8(0)]TJ T* 0 Tc (.1084/jem.20122252)Tj -1.7505 -1.1284 TD [(64.)-500.5(Zho)16(u)-469.8(BO,)-464.2(Yue)-463.7(R,)-468.5(Murp)16.1(hy)-463.8(MM,)-462.4(et)-473.6(al.)-461.4(Lep)11.8(ti)13.1(n-r)13.2(ece)17.4(pto)12.3(r-)]TJ 1.7505 -1.1223 TD [(ex)13.8(pre)11.3(ss)13.7(ing)-347.5(mes)15(en)13.8(chy)13.7(mal)-345.2(stro)16.9(ma)14.2(l)-359.4(cel)16.1(ls)-346.5(rep)11.3(re)11.5(sen)14.5(t)-353.3(the)-345.6(mai)14.7(n)]TJ 0 -1.1284 TD [(so)12.8(urc)17.4(e)-370.3(o)0(f)-362.5(bon)17.9(e)-370.3(for)13.2(me)14.2(d)-366.1(b)0(y)-366.2(a)0(d)13.8(ult)-359.1(bon)17.9(e)-370.3(mar)17.9(row)14(.)-366(Cell)-356.1(Ste)17.4(m)]TJ 0 -1.1223 TD [(Cel)15.6(l)-267.9(201)17.9(4;1)12.3(5\(2)15.5(\):1)16.1(54)17.9(168)17.9(;)-274(doi)18.4(:)-267.9(1)0(0)18(.101)17.8(6/)12.5(j)12.6(.st)13.3(em)14.2(.20)17.9(14.)17.9(06)18(.00)17.9(8)]TJ -1.7505 -1.1284 TD [(65.)-500.5(Kunisaki)-511.2(Y,)-514.1(Bruns)-513.2(I,)-514.9(Scheiermann)-516.7(C,)-511.2(et)-516.3(al.)-510.2(Arteriolar)]TJ 1.7505 -1.1223 TD [(niches)-648.2(maintain)-647.8(haematopoietic)-654.7(stem)-649.4(cell)-648.3(quiescence.)]TJ 0 -1.1284 TD .025 Tc [(Natu)-5.6(re)-1345.6(2)-5.6(013)-5.6(;5)-5.6(02\(7)-5.6(4).5(7)-5.6(3).5(\):)-5.1(637)-5.6(64)-5.6(3;)-1346.9(d)-5.6(o).5(i)-5.1(:)-1346.9(10)-30(.)-5.6(103)-5.6(8/)]TJ 0 -1.1223 TD 0 Tc (nature12612)Tj -1.7505 -1.1284 TD [(66.)-500.5(Tikhonova)-263(AN,)-271.9(Dolgalev)-266.2(I,)-270.9(Hu)-264.1(H,)-264.1(et)-272.3(al.)-266.3(The)-268.8(bone)-267.1(marrow)]TJ 1.7505 -1.1284 TD [(microenvironment)-1041.7(at)-1034.7(single-cell)-1044.1(resolution.)-1042.3(Nature)]TJ 0 -1.1223 TD [(2019;572\(7767\):E6;)-330.7(doi:)-334.9(10.1038/s41586-019-1394-x)]TJ -1.7505 -1.1284 TD [(67.)-500.5(Baccin)-359.3(C,)-352.7(Al-Sabah)-357.3(J,)-353.1(Velten)-357.4(L,)-355.9(et)-357.7(al.)-357.7(Combined)-356.8(single-)]TJ 1.7505 -1.1223 TD [(cell)-636.1(and)-632.9(spatial)-635(transcriptomics)-637.1(reveal)-637.4(the)-632.3(molecular,)]TJ 0 -1.1284 TD [(cellular)-349.8(and)-346.2(spatial)-348.4(bone)-346.3(marrow)-346.3(niche)-350.1(organization.)-350.5(Nat)]TJ 0 -1.1223 TD .0183 Tc [(Cell)-402.1(Bio)6(l)-402.1(202)6(0;22)6(\(1)6(\):3)6(84)6(8;)-402.1(do)6(i:)-402.1(10)-12.3(.10)6(38/s4)6(155)6(6-019-)]TJ 0 -1.1284 TD 0 Tc (0439-6)Tj -1.7505 -1.1223 TD [(68.)-500.5(Baryawno)-329.3(N,)-331.2(Przybylski)-326(D,)-331.2(Kowalczyk)-333.7(MS,)-325.8(et)-333.3(al.)-327.3(A)-331.1(cel-)]TJ 1.7505 -1.1284 TD [(lular)-262.3(taxonomy)-260.5(of)-264.9(the)-260.2(bone)-261(marrow)-260.9(stroma)-267.8(in)-262(homeostasis)]TJ 0 -1.1223 TD [(and)-273(leukem)10(ia.)-278.5(Cell)-270.7(2019;177\()15.2(7\):19151)15.2(932;)-274.5(doi:)-273.9(10.101)17.5(6/)]TJ 0 -1.1284 TD (j.cell.2019.04.040)Tj -1.7505 -1.1223 TD [(69.)-500.5(Ono)-727.8(N,)-727.6(Ono)-727.8(W,)-730.4(Mizoguchi)-733.6(T,)-727.9(et)-729.7(al.)-729.8(Vasculature-)]TJ 1.7505 -1.1284 TD [(associated)-546(cells)-543.8(expressing)-546.6(nestin)-545.9(in)-542.5(developing)-545.4(bones)]TJ 0 -1.1223 TD [(encompass)-469.2(early)-474.4(cells)-470.6(in)-469.3(the)-473.7(osteoblast)-469.9(and)-474.3(endothelial)]TJ 0 -1.1284 TD .0237 Tc [(l)5.8(i)-.3(neag)5.3(e.)-409.4(Dev)-409.5(C)6.6(el)5.8(l)-408.9(2)5.3(01)5.3(4;2)5.3(9)-.8(\(3\):)5.8(330)5.3(3)5.3(39)5.3(;)-408.9(d)-.8(o)5.3(i)-.3(:)-402.8(1)-.8(0)-19.1(.)-.8(1)5.3(0)-.8(1)5.3(6)-.8(/)5.8(j)]TJ 0 -1.1223 TD 0 Tc (.devcel.2014.03.014)Tj -1.7505 -1.1284 TD [(70.)-500.5(Chan)-320.6(CKF,)-320.8(Lindau)-323.6(P,)-320.3(Jiang)-326.7(W,)-321.7(et)-321.1(al.)-321.2(Clonal)-325.9(precursor)-326.9(of)]TJ 1.7505 -1.1223 TD [(bone,)-523.3(cartilage,)-525(and)-523.1(hematopoietic)-522.2(niche)-527(stromal)-523.5(cells.)]TJ /F9 1 Tf 8.9663 0 0 8.9663 59.7543 749.7069 Tm [(ORGANOID)-340.1(CULTURE)-336.6(DEVELOPMENT)-339.9(FOR)-334(SKELETAL)-332.4(SYSTEMS)-21292.4(555)]TJ ET endstream endobj 201 0 obj <>stream /GS1 gs BT /F6 1 Tf 9.2951 0 0 9.2951 82.9417 725.1588 Tm 0 0 0 rg 0 Tc 0 Tw [(Proc)-430.8(Natl)-432.2(Acad)-437.5(Sci)-427.8(U)-434.8(S)-430(A)-428.7(2013;110\(31\):1264312648;)]TJ 0 -1.1223 TD [(doi:)-334.9(10.1073/pnas.1310212110)]TJ -1.7505 -1.1284 TD [(71.)-500.5(Di)17(ng)-256.5(L,)-252.2(Sa)17(un)18(de)19.9(rs)-257.9(TL)14.2(,)-262.3(E)0(n)16.1(i)12.6(ko)18(lo)18.6(po)18(v)-262.4(G)0(,)-258(e)0(t)-260.1(a)0(l)20.5(.)-268.4(En)22.2(do)18(th)18.6(el)20.5(ia)14.4(l)-261.8(a)0(n)13.8(d)]TJ 1.7505 -1.1223 TD [(pe)19.9(ri)16.3(va)19.9(sc)14.7(ul)24.7(ar)-281.3(ce)15.7(l)12.6(l)0(s)-285.5(m)12.5(ai)20.5(nt)18.6(ai)14.4(n)-286.8(h)0(a)19.9(e)0(m)14.2(a)14(to)18.6(po)18(ie)14.4(t)12.6(i)0(c)-278.4(s)0(t)19.5(e)0(m)-284.6(c)0(e)21.8(l)0(l)19.2(s)0(.)-279.9(N)0(a)18.3(t)0(-)]TJ 0 -1.1283 TD [(ur)21.8(e)-272.7(2)0(0)18(1)0(2)18(;)12.6(48)18(1\()15.7(73)24.1(82)18(\):)16.3(45)18(7)24.1(46)18(2;)-262(do)18(i)12.6(:)-274(10)24.1(.)18.2(1)0(0)18(3)0(8)18(/)12.6(na)13.8(t)12.6(u)0(r)15.7(e)0(1)19.9(0)0(7)18(8)0(3)]TJ -1.7505 -1.1223 TD [(72.)-500.5(Sivaraj)-306.3(KK,)-302.4(Jeong)-302.9(HW,)-305.2(Dharmalingam)-309(B,)-303.9(et)-302.8(al.)-302.9(Regional)]TJ 1.7505 -1.1284 TD [(specialization)-330.4(and)-327.9(fate)-327.9(specication)-329.9(of)-325.9(bone)-321.9(stromal)-328.3(cells)]TJ 0 -1.1223 TD [(in)-579.1(skeletal)-578.4(development.)-580(Cell)-575.6(Rep)-576.6(2021;36\(2\):109352;)]TJ 0 -1.1284 TD [(doi:)-334.9(10.1016/j.celrep.2021.109352)]TJ -1.7505 -1.1223 TD [(73.)-500.5(Sacchetti)-385.3(B,)-377.1(Funari)-381.8(A,)-373.8(Michienzi)-383.5(S,)-381.3(et)-376(al.)-382.1(Self-renewing)]TJ 1.7505 -1.1284 TD [(osteoprogenitors)-280.3(in)-274.2(bone)-279.2(marrow)-279.2(sinusoids)-278.1(can)-277.2(organize)-277.8(a)]TJ 0 -1.1223 TD [(hematopoietic)-406.3(microenvironment.)-407.4(Cell)-398.8(2007;131\(2\):324)]TJ 0 -1.1284 TD [(336;)-335.5(doi:)-328.8(10.1016/j.cell.2007.08.025)]TJ -1.7505 -1.1284 TD [(74.)-500.5(Mabuchi)-672.9(Y,)-666.6(Morikawa)-670.9(S,)-667.9(Harada)-676.1(S,)-667.9(et)-668.7(al.)-668.8(LNGFR)]TJ 1.7505 -1.1223 TD (\()Tj /F11 1 Tf .3355 0 TD (+)Tj /F6 1 Tf .5489 0 TD (\)THY-1\()Tj /F11 1 Tf 3.5498 0 TD (+)Tj /F6 1 Tf .555 0 TD (\)VCAM-1\(hi)Tj /F11 1 Tf 5.2697 0 TD (+)Tj /F6 1 Tf .555 0 TD [(\))-417.2(cells)-421.8(reveal)-417.8(functionally)-424.9(dis-)]TJ -10.8139 -1.1284 TD [(tinct)-546(subpopulations)-545.6(in)-548.6(mesenchymal)-546.7(stem)-545.7(cells.)-543.9(Stem)]TJ 0 -1.1223 TD .0129 Tc [(Cell)-407.5(Rep)-5.5(o).6(rt)-5(s)-401.1(2)-5.5(01)-5.5(3;)-5(1\(2\))-7.9(:1)-5.5(52)-5.5(16)-5.5(5;)-407.5(doi)-5(:)-401.4(1)-5.5(0)-17.7(.).6(1)-5.5(016)-5.5(/j)-17.2(.)-5.5(s)1.4(t)-5(e)-3.6(mcr)]TJ 0 -1.1284 TD 0 Tc (.2013.06.001)Tj -1.7505 -1.1223 TD [(75.)-500.5(Buhring)-257.6(HJ,)-251.1(Battula)-256.5(VL,)-253.9(Treml)-258.2(S,)-253.2(et)-254(al.)-254.1(Novel)-256(markers)-256.7(for)]TJ 1.7505 -1.1284 TD [(the)-333.4(prospective)-331.7(isolation)-331.7(of)-332(human)-333.9(MSC.)-330.7(Ann)-331.4(N)-331.1(Y)-331.1(Acad)]TJ 0 -1.1223 TD [(Sci)-336.3(2007;1106:262271;)-330.3(doi:)-334.9(10.1196/annals.1392.000)]TJ -1.7505 -1.1284 TD [(76.)-500.5(Boiret)-377(N,)-367.7(Rapatel)-377.1(C,)-371(Veyrat-Masson)-377.5(R,)-377.1(et)-369.9(al.)-376(Character-)]TJ 1.7505 -1.1223 TD [(ization)-605.4(of)-606.4(nonexpanded)-605.9(mesenchymal)-607.7(progenitor)-600.8(cells)]TJ 0 -1.1284 TD [(from)-511(normal)-506.5(adult)-510(human)-510.7(bone)-511(marrow.)-511.1(Exp)-508.6(Hematol)]TJ 0 -1.1223 TD [(2005;33\(2\):219225;)-335.1(doi:)-334.9(10.1016/j.exphem.2004.11.001)]TJ -1.7505 -1.1284 TD [(77.)-500.5(As)23.4(l)12.6(a)0(n)-315.6(H)0(,)-312.9(Z)0(i)22.8(l)12.6(be)19.9(rm)22.2(a)14(n)-323.4(Y,)-319(K)16.6(a)0(n)19.9(d)0(e)19.9(l)-322.8(L,)-313.2(e)14(t)-328.9(a)14(l)0(.)-316.8(O)0(s)23.4(t)12.6(eo)19.9(ge)26(ni)24.7(c)-327.6(d)0(i)24.7(f)0(f)19.5(e)14(r-)]TJ 1.7505 -1.1223 TD -.0141 Tc [(en)-14.1(tiat)-14.1(i)5.1(o)-14.1(n)-587.7(o)-14.1(f)-596.1(n)-14.1(o)10(n)-14.1(c)11.9(u)-14.1(ltu)-14.1(r)7.7(ed)-605.9(immu)-14.1(n)10(o)-14.1(i)10.6(s)-14.1(olat)-14.1(e)6.4(d)-605.9(b)-14.1(o)10(n)-14.1(e)-591.9(ma)-14.1(r)9.6(r)-14.1(o)7.7(w)-14.1(-)]TJ 0 -1.1284 TD 0 Tc [(de)19.9(r)15.8(i)12.6(ve)19.9(d)-231.9(C)13.4(D1)22.5(05)24.1(\()]TJ /F11 1 Tf 6.3188 0 TD (+)Tj /F6 1 Tf .5428 0 TD -.0224 Tc [(\))-250.5(c)-22.4(el)-22.4(ls)-9.5(.)-254.2(S)-22.4(te)-8.4(m)-260(C)-9(e)-8.4(l)-22.4(ls)-253.4(2)-22.4(0)1.7(0)-22.4(6)1.7(;)-9.8(2)-22.4(4\()-22.4(7\))-22.4(:1)-22.4(72)-22.4(8)-22.4(17)-22.4(37)-22.4(;)]TJ -6.8616 -1.1223 TD 0 Tc [(do)24.1(i)12.6(:)-322.8(10)36.3(.1)24.2(63)24.1(4/)18.6(s)12.9(t)12.6(em)20.3(c)14(e)0(l)20.5(l)12.6(s)25.1(.)0(2)24.2(0)0(0)24.1(5)0(-)21.8(0)0(5)24.1(4)0(6)]TJ -1.7505 -1.1284 TD [(78.)-500.5(Battula)-561.4(VL,)-565(Treml)-557.1(S,)-564.2(Bareiss)-563.1(PM,)-557.6(et)-558.9(al.)-565.1(Isolation)-560.3(of)]TJ 1.7505 -1.1223 TD [(functionally)-290.7(distinct)-294.9(mesenchymal)-290.6(stem)-289.5(cell)-294.5(subsets)-288.7(using)]TJ 0 -1.1284 TD [(antibodies)-312.2(against)-318.5(CD56,)-312(CD271,)-312.1(and)-315.7(mesenchymal)-315(stem)]TJ 0 -1.1223 TD [(cell)-270.1(antige)10.2(n-1.)-265.1(Haemat)10.3(ologi)12.6(a)-278.8(2009;)12(94\(2\))12.9(:173)12(184;)-268.4(doi:)-273.9(10)]TJ 0 -1.1284 TD (.3324/haematol.13740)Tj -1.7505 -1.1223 TD [(79.)-500.5(Chan)-460.9(CKF,)-455(Seo)-458.9(EY,)-455.2(Chen)-460.9(JY,)-458.5(et)-455.3(al.)-461.4(Identication)-456.6(and)]TJ 1.7505 -1.1284 TD [(specication)-769.1(of)-771.1(the)-766.5(mouse)-772.1(skeletal)-767.5(stem)-771.4(cell.)-770.3(Cell)]TJ 0 -1.1223 TD [(2015;160\(12\):285298;)-335.5(doi:)-334.9(10.1016/j.cell.2014.12.002)]TJ -1.7505 -1.1284 TD [(80.)-500.5(Chan)-284(CKF,)-284.2(Gulati)-285.5(GS,)-279.3(Sinha)-281.7(R,)-285.6(et)-284.5(al.)-278.5(Identication)-285.8(of)-283.2(the)]TJ 1.7505 -1.1223 TD [(human)-272.9(skeletal)-273.4(stem)-277.4(cell.)-276.3(Cell)-276.8(2018;175\(1\):4)12.9(356.e1;)-273.1(doi:)]TJ 0 -1.1284 TD (10.1016/j.cell.2018.07.029)Tj -1.7505 -1.1223 TD [(81.)-500.5(Mizuhashi)-250.9(K,)-251.9(Ono)-252.1(W,)-248.5(Matsushita)-255.7(Y,)-245.8(et)-254(al.)-248(Resting)-251.9(zone)-253(of)]TJ 1.7505 -1.1284 TD [(the)-424.9(growth)-425(plate)-422.7(houses)-424.2(a)-425.2(unique)-425.4(class)-427.6(of)-423.5(skeletal)-425.9(stem)]TJ 0 -1.1223 TD .025 Tc [(cell)-5.1(s)1.4(.)-450.8(N)-1.1(atu)-5.6(r)-1.9(e)-449(2).5(0)-5.6(18;)-5.1(56)-5.6(3\(77)-5.6(30)-5.6(\):25)-5.6(4)-5.6(258)-5.6(;)-444.2(d)-5.6(oi)-5.1(:)-444.2(1)-5.6(0)-30(.10)-5.6(38/)]TJ 0 -1.1284 TD 0 Tc (s41586-018-0662-5)Tj -1.7505 -1.1223 TD [(82.)-500.5(Ambrosi)-350.7(TH,)-351.5(Scialdone)-357.2(A,)-349.5(Graja)-354(A,)-349.4(et)-351.6(al.)-351.6(Adipocyte)-351.7(ac-)]TJ 1.7505 -1.1284 TD [(cumulation)-337.1(in)-341.3(the)-333.4(bone)-340.2(marrow)-340.2(during)-338.1(obesity)-338.6(and)-334(aging)]TJ T* [(impairs)-346.6(stem)-344.4(cell-based)-347.9(hematopoietic)-345.4(and)-346.2(bone)-340.2(regener-)]TJ 0 -1.1223 TD .0241 Tc [(ation.)-409(Cell)-408.5(Stem)-414.7(Cell)-408.5(20)5.7(17;20)5.7(\(6\):771)5.7(784)5.7(.e6)5.7(;)-408.5(doi:)-408.5(1)-.4(0)]TJ 0 -1.1284 TD 0 Tc (.1016/j.stem.2017.02.009)Tj -1.7505 -1.1223 TD [(83.)-500.5(Ambrosi)-545.9(TH,)-540.6(Sinha)-544(R,)-541.7(Steininger)-544(HM,)-537.9(et)-540.6(al.)-546.8(Distinct)]TJ 1.7505 -1.1284 TD [(skeletal)-523.5(stem)-515.2(cell)-520.2(types)-521.9(orchestrate)-521.2(long)-518.4(bone)-517.1(skeleto-)]TJ 0 -1.1223 TD [(genesis.)-336.5(Elife)-337.3(2021;10:e66063;)-334(doi:)-334.9(10.7554/eLife.66063)]TJ -1.7505 -1.1284 TD [(84.)-500.5(Debnath)-303.2(S,)-295.9(Yallowitz)-303.3(AR,)-293.4(McCormick)-301.7(J,)-298.2(et)-302.8(al.)-296.8(Discovery)]TJ 1.7505 -1.1223 TD [(of)-277.1(a)-278.8(periosteal)-282.2(stem)-277.4(cell)-282.3(mediating)-282(intramembranous)-278.8(bone)]TJ 0 -1.1284 TD .0251 Tc [(form)-5.1(ati)-5(on.)-566.6(N)-7.1(a)2.5(t)-5(u).6(re)-570.9(201)-5.5(8;)-5(562)-5.5(\(77)-5.5(25\):)-5(1).6(3)-5.5(31)-5.5(39;)-566.1(d)-5.5(o).6(i)-5(:)-566.1(10)]TJ 0 -1.1223 TD 0 Tc (.1038/s41586-018-0554-8)Tj -1.7505 -1.1284 TD [(85.)-500.5(Duchamp)-285(de)-285.1(Lageneste)-286.9(O,)-282.4(Julien)-283.6(A,)-282.4(Abou-Khalil)-284(R,)-279.5(et)-284.5(al.)]TJ 1.7505 -1.1223 TD [(Periosteum)-476.1(contains)-471.1(skeletal)-474.7(stem)-472.5(cells)-476.7(with)-470.6(high)-469.6(bone)]TJ 0 -1.1284 TD [(regenerative)-398.3(potential)-397.8(controlled)-400.8(by)-396.7(Periostin.)-392.4(Nat)-402.1(Com-)]TJ 0 -1.1223 TD [(mun)-335.5(2018;9\(1\):773;)-334.3(doi:)-334.9(10.1038/s41467-018-03124-z)]TJ -1.7505 -1.1284 TD [(86.)-500.5(Ortinau)-593.5(LC,)-586.5(Wang)-594.7(H,)-587.3(Lei)-591.5(K,)-593.4(et)-589.4(al.)-589.5(Identication)-596.9(of)]TJ 1.7505 -1.1223 TD [(functionally)-345.6(distinct)-337.6(Mx1)]TJ /F11 1 Tf 10.3748 0 TD (+)Tj /F6 1 Tf .555 0 TD (alphaSMA)Tj /F11 1 Tf 4.3305 0 TD (+)Tj /F6 1 Tf .8966 0 TD [(periosteal)-343.1(skeletal)]TJ -16.1569 -1.1284 TD [(stem)-265.2(cells)11.2(.)-274.5(Cell)-270.7(Ste)11.3(m)-274.2(Cell)-270.7(2019;)12(25\(6)15.4(\):784)15.7(796.e)13.4(5;)-274.2(doi:)-267.8(10)]TJ 0 -1.1223 TD (.1016/j.stem.2019.11.003)Tj 25.2569 70.8975 TD [(87.)-500.5(Tevlin)-518.1(R,)-517.3(Seo)-519.9(EY,)-516.2(Marecic)-519(O,)-520.2(et)-516.3(al.)-516.3(Pharmacological)]TJ 1.7505 -1.1223 TD [(rescue)-484.6(of)-472.2(diabetic)-481.7(skeletal)-480.8(stem)-472.5(cell)-483.6(niches.)-477.5(Sci)-476.6(Transl)]TJ 0 -1.1284 TD .0185 Tc [(M)-5.3(e)2(d)-402.5(2017;)-5.5(9\(372\):eaag2809;)-408(doi:)-408(1).1(0)-18.2(.1126/scitranslmed)]TJ 0 -1.1223 TD 0 Tc (.aag2809)Tj -1.7505 -1.1283 TD [(88.)-500.5(Jones)-332.5(RE,)-324.2(Salhotra)-330.8(A,)-325.1(Robertson)-328.3(KS,)-328.1(et)-327.2(al.)-327.2(Skeletal)-326(stem)]TJ 1.7505 -1.1223 TD [(cell-Schwann)-280.4(cell)-276.2(circuitry)-274.9(in)-274.2(mandibular)-279(repair.)-275.7(Cell)-270.7(Rep)]TJ 0 -1.1284 TD [(2019;28\(11\):27572766.e5;)-346.2(doi:)-341(10.1016/j.celrep.2019.08)]TJ 0 -1.1223 TD (.021)Tj -1.7505 -1.1284 TD [(89.)-500.5(Chan)-436.5(CKF,)-430.6(Chen)-430.4(CC,)-430.8(Luppen)-433.9(CA,)-433.7(et)-430.9(al.)-430.9(Endochondral)]TJ 1.7505 -1.1223 TD [(ossication)-296.1(is)-285.5(required)-288.1(for)-291.7(haematopoietic)-288.7(stem-cell)-293.8(niche)]TJ 0 -1.1284 TD .0254 Tc [(fo)-5.2(rmat)-4.7(io)-5.2(n.)-566.3(Natu)-5.2(re)-564.5(20)-5.2(09)-5.2(;45)-5.2(7).9(\(7)-5.2(228)-5.2(\):4)-5.2(90)-5.2(494)-5.2(;)-565.8(d).9(o)-5.2(i)1.4(:)-565.8(1).9(0)]TJ 0 -1.1223 TD 0 Tc (.1038/nature07547)Tj -1.7505 -1.1284 TD [(90.)-500.5(Street)-318.3(J,)-310.4(Bao)-308.3(M,)-316.6(deGuzman)-313.8(L,)-307.1(et)-315(al.)-309(Vascular)-315.1(endothelial)]TJ 1.7505 -1.1284 TD [(growth)-498.2(factor)-501.3(stimulates)-499.4(bone)-498.8(repair)-495.2(by)-500.4(promoting)-496.1(an-)]TJ 0 -1.1223 TD [(giogenesis)-330.2(and)-327.9(bone)-328(turnover.)-326.7(Proc)-327.2(Natl)-328.5(Acad)-327.7(Sci)-324.1(U)-325(S)-326.3(A)]TJ 0 -1.1284 TD [(2002;99\(15\):96569661;)-335.5(doi:)-334.9(10.1073/pnas.152324099)]TJ -1.7505 -1.1223 TD [(91.)-500.5(Ambrosi)-295.8(TH,)-284.4(Marecic)-299.4(O,)-288.5(McArdle)-292.6(A,)-288.5(et)-290.6(al.)-290.7(Aged)-286.9(skeletal)]TJ 1.7505 -1.1284 TD [(stem)-405.4(cells)-403.5(generate)-410.1(an)-400.9(inammatory)-403.6(degenerative)-408.2(niche.)]TJ 0 -1.1223 TD .0124 Tc [(N)-7.6(a)-4.1(tu)-6(re)-400.6(2)-6(0).1(2)-6(1).1(;)-5.5(597)-6(\(78)-6(7).1(5)-6(\))-2.3(:2)-6(56)-6(26)-6(2;)-408(doi)-5.5(:)-401.9(1)-6(0)-18.2(.).1(1)-6(038)-6(/s)-5.2(41)-6(58)-6(6-)]TJ 0 -1.1284 TD 0 Tc (021-03795-7)Tj -1.7505 -1.1223 TD [(92.)-500.5(Engler)-435.7(AJ,)-428(Sen)-434.5(S,)-430.1(Sweeney)-432.9(HL,)-430.8(et)-430.9(al.)-430.9(Matrix)-432.4(elasticity)]TJ 1.7505 -1.1284 TD 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