Studies of the development and decay of the human frame

394 Pages·1968·33.805 MB·English

by  Josep Trueta

'ForL itofhefae l Fll esh
itsh Bel otohde "r eof
(LeviXtViIcIu)s
TO AMELIA, MY LIFE COMPANION
Studioefts h eD evelopment
andD ecaoyf t heH umanF rame
JosepThr uetaF RCSF ACS
MD
Nuf{ielPdr ofessEomre ritoufsO rthopredSiucr gerUyn,i versoifOt xyf ord.
DoctoHro noriCsa usaU niversiotfOi xefso rdB;u enoAsi resR;i od eJ aneirBoo;g otaG;o theburg.
HonoraPrrye sidoefnt th eS ocieItnet ernatidoenC ahlier urOgriteh opedieqtTu rea umatologie.
HonoraFreyl loowft heA mericaCno lleogfeS urgeonRso;y aClo lleogfeS urgeoonfsC anadaB;r itiOsrht hopredAiscs ociation
{1940R)o;y aFla culotfyP hysiciaanndsS urgeonGsl,a sgowA;m ericaAns sociatfioortn h eS urgeroyf T raumaW;o rcester
CollegOex,f ord.
HonorarMye mbero ft heS ocieFtrea m;aids'eO rthopeedtTi rea umatologSioec;i elttaa lidainO ar topedei Tar aumatologia;
DeutschOer thopadisGcehsee llschSaofcti;e tOarst hoprediSccaa ndinaviScoac;i edaLda tino-Ameridcea Onrat opedyi a
TraumatoloSgolcai;e daBdrea si/deeiO rrat opeed iTar aumatoloSgoicai;e dCaodl ombiadneCa i rugOfrat opedyi cTar aumatologia;
AcademidaeM edicidneaB uenoAsi resS;o ciedCaadt aladneaP ediatrfa.
CorrespondMienmgb ero ft heA mericAanc ademoyf O rthopedSiucr gerlyn;s tidt'Eu stt udiCast alanSso;c iedEasdp anodlea
CirugOlrat hopedyi Tcraa umatoloAgclaad;e midee ChirurgPiaer idse.
Of{iciedre l aL egiodn' HonneuCro.m endaddoarO rdedno C ruzeirdoo S ulB,r azil.
WILLIAM HEINEMANN MEDICAL BOOKS LTD.
LON DON
FIRSPTU BLISH1E9D6 8
©JOSEPTHR UET1A9 68
PrinitnGe rdeB arti attta hiPeni tmParneB sast.h
PREFACE
THIbSo odke awlistt hhfa es cinsattoiornftyg h d ee veloopftm hehenu tm an sktehflauetnt doanm,ie nnntseatrlr ucture
whipclha mcaensa boavleol t hleirv cirnega tfuoriret sm ,ai isn dluyte o uintisq ue rtehfamitan ne mscetanantn d
ereacnttd h ahtib sr aipnl aichsei dga hb oavleol t her organsf.o trNh peirvsie vrithlhepee aglydeese,s a srb,lo yt,h
ins uffearniidnnt g h leo osfms o biilnti htleya sttea ogfle is fe.
Asa no rtrehdoispcu rgfeoorn 4o0vy eearIr h sa,vw ei tnde-sasneedv ecno ntritbou-ttcehheda ngaitntgio tfu de
thmee mbeorfms ys peciwahlofri,ot may c arpeanptprryot aooc rht hophreadviec sat,g a rkoewnii nntge irnte hset
stuodfyt hnea tuarnerd e aocntosif t hlei vtiinsgsw uheicsco hn sttihtseuk teel eIttio stnh .ii nst ewrheiscth has
increaospiennugeptld hy le a borfaatlcoiirttyiou e sas,n hda nso otn leyn laoruguren dd erstoaftn hpdehi negn omena
inveodil ntv hper ocoefgs rso watnhdd i seoafts hehe u mafrna meb,u t ahlasmsoa diept o ssfiobtrlh oeer tphreodic
surgteoco onn trtiosb yumtpeoa sncidoa n feroenpn ucreresel sye saurjbcehc ts.
Persosnianlmclyeey a ,rm leyd idcaayilsn1 2 96w heInw adsi crteetdoi nvesetxipgeartiemw eanyotsfpa rloldyu cing
joiinntt hsfee m orsahflat o fd ogIsh ,a vnee vleorms yti nteirnte hssett uodfty i sbseuhea vainoi unrt,ew rheiscth
sermveew de dlulr tihnSgep anCiisvWhia lrT .h eIrw ea asb tloed evealb oiopog limceatlh ooftd r eatiwnogu nwdasr
anfrda ctsuw,rh eiwcahrs e spofnosmriy bc loem itnolg i ivneB ritian1i 39n9 W.h iilnte h ciosu ntthrreye ,s earch
faciwlhiitwcieher asev ailtaob mlmaeed peo sstihbel eb roamdyef niieonlfagd c tosifoot nh aIwt a ss ooanb lteo
contrtiotb huietn ev estigtaheteffi eoocnfit m omno biluipzoatnth lieyo mnp hcaitriccu altat thWieio lnl iam Dunn
SchooofPl a thowlhoigIcyj h,o idn sehtolrbye foPrreeos fsHo.rF lroeayn dD rC.h awienr teoe mbaornkt heir
reseoanrp cehn icwihlihlcaihcsn h antghehedit soroyfm edicMiync eo.n cientr hnce i rcuolfat thbielo onho add
incredausretidhnw gea irnC atalwohneiIrah e,a c do elcltdeadto ant hdee cirsoiolvftee h cei rcuilnma utsicoalnne ds
bonien htehael oifwn ogu nTdosg.e twhietDrhr J .o hBna rnIew sa,sa blteoi nveastttehi eegff eocfst houcpko n
thvee ssoeftl hsee x tremaintdli aetwsei,rtt hhfr ei encdollyl aboofrD art(.il oaPntr eorfe sKseonrn)Fe rtahn klin,
DrA.l frBeadr calnaMdyi sMsaj rorPirei chsaorojdno, i bnyeD drP .e tDearn iIse tlu,d tihreeed n cailr ciuolnIa.ntt his
worwked evelaom peetdho ofvd a scvuilsaura lbiymz iactrriaoodni ogforraw phhiDycr,hB. a rcwlaamsya inrleys pon­
sibwlhei,cw hh,eI nu seidt coijnnu ncwtiittohhn ao tfS paltteoho obltttazii snt sruaen spgaarvedene ctyao,int l hse
intraocsisrecouuwlsha ithciahond no bte eanv ailbaebfolrIeem .u spta y ttroim byfu rtiePe rnodfe sAsmoorr ofosro
three finemoeftn hStets a lteholozf w theiwccehhw n eirtqeomu aek,ce o nsiduesrieant b hcleoe u orfet shfo el lowing
twenyteya rs.
Witthhp eu bliocfa tboiouoornSk t udoifte hsRe e nCailr cuilan1t4 9i7mo,yn w oroknt hkei dneyt oac naa bmreu pt
endT.h iwsa sd uteo ampyp oinatsNm ueffinetPl rdo fesosfOo rrt hopSreudrigacet tr hyUe n viersoifOt xyfo rd.
Howevtehrpe,u blicoaft iowoounrro k nt hree ncailr cuwlaassto iononon t ibcyse odm oer thopsreudrigcew ohnos
suspetchtatethdm e e thoofdv sa scvuilsaura lwiezh aatddei socnr ciobueldd b eua slestdoos tudyi ntthrea osseous
circounlA.as et airals1y 49 8D rR..d eM arnee,ff frBoeml giinuqmu,ii nrttehodpe o ssiobfui sliiontugmyr e thtood s
investthivega astceu loaft ubroTenhb eeo. ohke pubiln1i 9sw5ha1eas dn e xceclolnetnrti tbotu htseiu obnjw ehcitc,h
morree centh acwsoo inrrfkm ienid tm sa ilni nes.
Duritnhlgea tswte ynteyat rhasem ouonfit n formtahthaiatbos en e gna thieosrf se du cmha gniptaurdtei,ci unl arly
thlea dsetc atdheIa,t t h iinjtksu titfioae tdt etmoop rtg aans iuzsbet anatmioauolnft t h aev aimlaatbelirenai l aolg ical
sequencew.h alTht ha ivaset tiesmt podt owe idt thhm ea tecroilallie nct thleead b oraatnowdra iroedfss the Nuffield
OrthopCreednitacrsw e e,al slt hweo rokfo thienrv esttihgoauItmg ouhrs sct,o nfetshsta htbi oso iksf afrro m being
comprehensive.
Myi ntenitnsio totopn r odauncoet theebxrot o akst hfei eilasdl reeaxdcye lcloevnetbrlueystd i ,m ptlody e scribe
thseu cceosfse ivoennw thsi cihnt, h ceo urosfte i mhea,rs a itshejdee lolryg anoifst mhssee ian t hpea leo­
litdhaiwocnf Etahrett oth h ceo mpaltiescdt ruoctftu hrreie gi indnfr earm oefp resenmta-nd.a y
Top revaef nete loiffrn ugs trIam tuiswotan rt nhr ee adpearr,t icihufel i aasr r elsye waorrckhet rhf eii eonlf od s teo­
genetshimasat n y opfo itmnhatedsi e nt hepsaega ersse t uinldlde irs cuasnsidin o no,ct ahseers i st hnbeomyr e ea ns
unaniamgoruese wmietmnhytc onscilounHosw.e vaesIr h ,a vael resaadimydy , intiestno ct rieoaans t eeq uence
whiicsfhu ,c cespsrfeusleglniytva eev dsi,s iumaalog fte h fien arle souftl htma atg nificcreenaittm ipvuel ses hwahpiecdh
thien itial enteor Ggieynn earsse"i flsei rgirhnetttd"ho ca otn glaotomefeh ropaen frdu straticoonn swthtiihtceuh t es
human bAesai m negd.i mcaanla, n adl saosa l aborwaotrokrIeyh r a,va ed eeapd mirfoart iottnhw efo u tlilm e
spelciistathcsel, i niacntid hasnec, i eanmtoinswgth ,oI mh avoen lbye eanp art tHiomweervi.emt ra ,yw eblelt hat
myt wsoi diendt ehrapeses rtm imtett oae tdt eamnep ntt erwphriicshe a ed iedtihcesarct ieeodnr atr iesptuc tleidn ician
wouhladv foeu nlde isnsc ltioun nedde rtake.
JT.R UETA
ACKNOWLEDGEMENTS
To pay rdeuceo gntio ttiaholonlws heo hcaovnietb rutteom da kteh bioso pko sswiobulrleed,q uainer neo rmously
lengltiohsnfyta mweistt hhe ceorfst tabiielnilitn nygc ompIlnte hpteae g.te foso llonwa,m oetfsh weto hrek wehreosi ther
aloonrwe i tmhyc ollabohraavpteri oovnmi,do esodtf mtahtee forrit ahbleo oakr deu lrye poarnttdeo td h egmo es,
witmhyt hantkhssew, e meetm oroifte hsre e ceddaiynwsgh ewnew orkteodg etMhyeo rla.ds sisatnadn tssc holars,
mosotfw hoamr neo wl eadoeftr hssep ectiyia ntl hieoiwrcn o untsrhioeucslo,dn stihdbieosro a ksa h umbhlomea ge
tot hewiorr k.
Howevseormn,ea mepse oowfph loeh ave dbiereecnco tnlcyew rinttehhdt ec e hnipcraold uocftt hibioosno nke ed
tob em entitoomn aekdpe u bmlyir ce cognn.Ti htfeii oirDssrt K. e thuLriatthtow l heo Im o wteh oer iginal electron
microysp chooptogwrhaipschehsr tvoes dt utdhyte o pogroafp ohtsyht ee ogceenlaelttts hit eci mweew erien terested
int hree latioofn svthehisewps a elalln tdh oes teoSchyheta eas.l spor ovimdewe idta hn umboefer x cellilgehntt
microspchoopteo garnahdpa hhsse lpientd h e sotftu hdceyh emiosfct arlycc aitfiiIo anm.h apptyob ea blteo
thahnekpr u bliTcoDl ryP..e tBeurl loaunDgdrh H ..E zra-ICl oihtknoee x pregsrsa tmfiyot truh dee thhegelaypv e
mei nt hfiee lodfts h eriers peaccttiivtvehifi etr yas,sta h isgtioaslntod stehceoi ntndh e ex perwiimtiehmn ptl iann ts
transcphaarmebnietnrr asb beiatrT'sos.M issL iMt.c hMfriD.e. l dCW,h. a ralneMdsi sGs.S toctkeecsh,n iinc ians
the DepIag rrtaumtleealnfcytk nowtlheecdiogrne t roinbsMu.rt D.i. D ruarnyMd r . ERm.m antuoeomlka noyft he
photogarnatdph Dhees p artomfRe andti oolfto hgNey. O.uCn.dt ehrde i reocftD irFo..nK empa nJd.B o ldwearso
responsniebaalrlell ycf lotirhn reia cdailo gFrianpahlml sul.sye t,x pregsrsa tmitytot u hdopese eo wphlwoee rpea rticu­
larcloyn cewrinttehhdl e i teprraordyu ocftt ihoen o tfte hxbeto o.kF irMsits,s Joans omG aensysye, a mryfos r
perssoencarlea tnatdrh ayed minisotfmr yDa etpoar rtwmheonustne,t ierffionrgt s and tsakssilokml u cmha de my
lighStretra.. M aFrlooatrnoMad r sJ.J. o hndseoawnli ttt hht ey poifnm go sotft htee xItm .u stth aMnrksE .l isabeth
Gilwih,op atiernetaaldlty lh tee axtn mda dmea ncyo nstrsuucgtgievsaetn iadom nesn dmeMnrtW.s. M. . S teel,
FR..C.cSl.o,ts helesi osftt hpee opwlheoh avmea dteh peu blicoaftt hsibi oono pko ssiAbstl hee. olfta hset
UneirvsLietcyt uurnedrmeserh eh abde einnc hragoef utnhree watradsoikfns gu pertvhipseri ondgu cotfti hoen
boomko,so tfw hiictsh obe donem yia nb senHcewe h.oh aadl rewaodryk ed ionnt rtahocesi srecouublsea fotrieo n
joinmiynD ge partwmaepsna tr ticulartloy meaan skdueoc wceedhs ists a osafkn Idg rateafcuklnloyw hliesd ge
contribution.
Thpeu bliosfth heer sbM oeosksW,ri sl.lH ieaimn emadnincB aoMloe khsa,vp el aacletldhr ee qufiaircleidat tmi ye s
disptooms aaklte h beo orke adaanbdl ep lteola osoaaktnT .to t hem Iaa dldsrome yts hsa nks.
J.F .
ERRATA
AcknowledgeFmoernJ t.F s.r eaJd.T .
Page3 0 4tpha ragrfoarpF hi g3s3.a nd3 4r eaFdi g2s6.2a nd2 63.
Page6 5 Leegdn toF ig8.2 s hourleda Kda ryotoyfpa e p atiweintth
chromatin-ngeognaatdiaavplel asZina+ I = 45 (XO).
TurneSry'nsd rome.
Pag1e0 2F ig1.63 i nl egednedl e(t 6e0 a)n d( 320).
x x
Pag2e3 6P enultilmiandteee l e(tFei 3g3.0 ).
Pag2e6 3 Fig35.2 s houbledF ig35.3 .
Pag2e64 Fig35.3 s houbledF ig35.2 .
Pag2e8 9F ircsotl umlna spta ragrfoarpd hi slocraetaiidon nf ection.
Pag2e97 Secocnodl um3nr ldi nfoer F ig.r e2a3Fd i g1.8 8.
Pag3e63 Fig50.3 s hourleda Fdi g50.4 ( aa)n d( b).
PARTI HISTORICAAPLP RAISAL
THROUGHOUtT he tahgoeer si,og fbi onn e has been-and
FROM GELT OB ONE stii-sloln oef t hmeo sfta scipnraotbilfneagmc nsio ntg
only biboulatol gspiohs iotlpsoh searnsed v etnh eolo­
gians. tHhosewc eiveesnrtt,ui odffbyi o cn beo,ta hsa
tisasnudae s a no rgadnin,do bte gin Coumnpttioln
Haveirn1s 96 1d escrtihcbeae ndia nlb so nweh ibceha r
hinsa meT.h en exitm porctoannttr iwbaumsta idoen
byS tepHhaeln(e 71s2 7w)h,eh nep ublitshehexe pde ri­
menwti tbho nmea rkiwnhgidsce hm onsttrhaattthe ed
CHAPTER1 lonbgo ngerso iwn leonnglatytth h eeixrt irteimes.
Sooanf terwfaorldlsot wheoed tu stanwdoirnokgf
HenrDyu hamdeeMl o nceIanu1 .47 2h ep ostulated
thatthg er owotfbh o nwea ssi mitlota hrao tfa t ree
andc onsitdhearbteo dnf eo rmadteipoenn odnet dh e
perio;si tnhe iuosmw nw ord'sa,s e xaong enous stem
OSTEOGENESIS
grofrwosm t hien nleary oeftr h bea kr'I.np lacai ng
cirocfls ei lwvierrse u bperiaogsatitinhascelto l ryt ex
ofa l iving heap neirmfoartlmh,eee dx perwihmiecnth ,
evemno rteh atnh aotfH almeas,yb ec onsitdheer ed
basoifes x periomsetneto;asl luo bgsyeqhueefo nutnldy ,
threi ng emibnteh sdeahd feotdft hbeo naen d concluded
thatthl eo nbgo ngerso iwnt hickbnyae pspso sition.
Unfotrunatheetl hyo,u tghhtitos tbheoe n lgye nesis
ofb oneD.u hamtehle'owsra ysk e patl idvuer itnhge
Fi1g . XIXtchen twuirttyhh s eu ppoofmr etna sd istinguished
SpecifmreonJmso hHnun tewro'rsok n asS ym(e1 4 80)F,l our(8eI4n 2sa) n Odl l(i81e6r7a )n d
bonger owtphr eseravtte hdeR oyal irte maainlemudon scth alulnetMniaglce edwp eunb lished
ColloefgS eug reonEsn,g la(nRde.p ro­
himsa steripn1i 9e,1ic 2new hicphr,o bafobrtl hye first
ducedb yk inpde rimssoifot nh Pe resi­
denatn dC ounocfit lh Reo yCalo llege timaes,e pptriocc ewdeurreeem sp loiynee xdp erimental
ofS ugreoonfsE ngland.) srugery.
Contrtaotr hyoe p inoifDo unah meAll,b recht von
Hall(e71r6 3a)n dJ ohHnu nt(e81r3 5a)t triab uted
promipnaerntt ipnr octoehfsbe so n foer mattioo n
Fi1g . thvee ssIetwl assa.m isfortthuanntee itohfte hre m
seetmosh avues eedx perimmeetnhtoatdlocs o llect
evidienns ucpep ooftr htev iire wtsh veoa ns courliagri n
ofb onTeh.ii assl lm otrhteeo lbaem enctoends idering
thabto tHha llaenrdH untweerr eex celelxepnetr i­
mentaalsci assntt sbi,els le einnt hcea soefH untbeyr
examitnwioon fgt hsep eciomfhe inwsso rokn b one
growth partte hsee rvRCeoodyl alolef Sg uer geoofn s
Engl(aFnid1)g ..E nchonodsrsailfi ciangt rioowni ng
bonesa pwpaasr neonmtte lnyt iuonnte1id38l 6w hen
Miescmhaedrte hf ei srustbtas nrteiefraeltn oct eh is basic
mechaonfti hsgemr owptrho cTehseesa .r lcioemsptl ete
accoouftn htge r owptlhaw taets h oaftT odadn Bdo w­
man( 8145w)h icwha ss oofno llobwyte hdpe a ins­
takisntguo dfSy h arpey, iantp hpfieef aterhdi intgi on
ofQ uaiEnel'mse notfA sn taom(y81 48t)h,e naontde s
refereonfcL eesi doyfP hilad(e1l8p4ah9ni)dat he
,. extensstiuvodefyT omeasn dD eM ogran( 8153).
.. Howeviewtra ,ns o utn t1i58l8w ,h eMnii lpluebrl ished
r' ..., • .. /, • ,./ -' his cnloawsw soirockn t hmee chanoifes pmi physeal
'.1,,. • ; .., • .. # , ,F/,/,. '1i:. cartciallaicgciieaof ttnh,a tthf ei fruslt l aocftc hoeu nt
basic meocfgh raonwwitasphsmu bliIsnth hetedw .e nty
yeaforlsl owtihnpegu blicoafMt uilolnew ro'rstk h,e
.fr.;1: /•'if detaainlaetdao nmfdyu ncotfti hoeenp iphcyasretaill age
.,,. l- 'H •;.•, I•
•,It� )":.1·11..: .. -.:/ becanmeea ralswy e lkln owanst heayr teo daIyn.
somree spseucctahsst , h orseeefr rtiotn hgp ea rt played
byt hvea sculaadtjuatrcoeet nhetep iphcyasretaill age,
OSTEEONGSEIS 3
somaeu thoofpr assg te neratitoohn asvc eos leleemc tiendtt ohm ea rrsopwa creest,a iiwntii nttghh ivena scular
morien formtahtaiinsou ns uatlol yfob uen idn thleu meansi, dt o ewsi tthh e sfailnsieun sep eonfps airo­n
text-obfot ookdsaT yh.e main reiapssr oonb ­fotri cutlsheeiisds np erfumseitohnop drsit oomr i croradio­
abltyh rea riitnpy r,e sdeanyst t udoifte hsue,s oef grapahnyod t hperro cedures.
vascuplearrf ubseifoorsnee ctifoornm iincgr oscopic
examinaat teicohnn,wi hqiuwceah ws e lkln owtnot he
THEN ATURAEN DO RIGIONF T HE
expeorftt hsne i netceeennttuhr y.
CHONDROBLAASNTD O STEOBLAST
A remarksatbefolpre w ardt hwfiear srs etfe retnoc e
Theg rowmtaht,u raatniddo enc aoyft hheu man
thbeo ne-forcmeilbnlyGgs o od(s81i4r5 fo)l,l obwye d
frammea yb ea scriinbt ehfiden aaln altyost ihbsee ­
Virchdoews'csr (i81p5t3oi)fto hnoe s teoorci adl cifiable
haviooftu wroc eltlhsce,h ondrobtlhaoess tt eaon­d
matreilxa bobryat theoeds teogceenleIltnsi1 .c68 4
blabsottr,he lated omfte hmmebe esresn cfahmyimlayl
Gegenbcaouirnt ehndeo wu nivearcscaelplntyae mde
oft iscseulel s.
ofo steoabnldfia nsatlM layc ew(e11n9)2 d emolished
wharte maionfte hdDe u hamtehle oorfty he ex clusive­
nesosft hpee rioasstb eounfmeo rmtiinsgs ue. The Chondroblast
Meanlwehg,ir eiantt ehraebdse tes nh oiwtnnh oer iginT heo rigainndc haractoeftr hicesh toincdsr oblast
oft hoes teoabnlidait shs ta rpdolsys ifibnladew rtiot earr bee ttkenro wtnh atnh oosfet hoes teoabnlda st
ont hseu bjoefoc stt eowlhoodg oyen so ati hri vsi ewsc onuseenqtlleyss suj bcett od ebaIttei .sr ecognized
ont hmea ttSeorm.ae u thwoerrosef o pintihoatnth e thatthc eh ondrroebslufralostmts tadhaep tatai on of
osteodbelraisfrvtoe mcd a rticella(lgKseo ll1i58k3;e r,m esencthiysmcsaeutlleo cl e rtcaoinnd iatcicoonmsp any­
Miil1l58e8;rW ,a lde1y68e5;rG ,e genb1a68u7ar,n, d inlgo caanlre mia.c hoTnhder coacsnyu trev ilvaec kt he
others). Sebveelriatelhv atewthdoo e sr tkeeorgse noefbt lioco d infli otcwsl ovsiec iwniitthylo oustii ntgs
celolr igifnraotwmeh di btleo ocde l(lWso l1ff78,0; reprcotdicuvaep acpirtoyv,i ded tihnatte rrmeigtutleanrt
Scahffer1,88 8; Kazzan1d09e0;r J,e rusa10l9e1;m ,c omprehsesltiphospen r ogreosfts iisosnu et oafl nudi ds
andre ceDnatnlaiyns Dd o zin16e95l),H. o wevtehre, from tcheelT lh.ea daptoatfti hocines tl ola nremic
most widely hbeeletdnh abte olstitheeefo bhlcaaossn td iitssiu octnhhs aa r ti ch flbolwio niot ndse ighbour­
occuarsas m etapolfam seisae nchoyrcm oanln ecthiovoemd a ys erioiumsplaiyit rss u rviAvnaale.r obic
tiscseuleAl mso.n tgh lea rnguem boefsr u ppoorft egrlsy coallylstoihwcsesh ondrtoosb ulravasintmvd ue l tiply
thviise own layfe ww iblelm entihoenresedu, ca hs int haeb seonfoc xey gen.
Rena(1u 88t6 B)u,n t(i10n96gK) l,o (1t 9z) 1a 6nMda xoiwm Oneo ft hmeo sitm porcthaanrta ctoefrt ihset ics
anBdl oo(9m15 7L)e.r iacnhPdeo li(c1a9rt2dh6 o)u ghcth onydtrieosi c tasb iltiort eyp roudnudcpeerr e ssures
thaots teobwlearsesi tmspu lnes peficbirfiocc ayntde sw,h iwcohu ilndh itbhiet reproduocfto itvhee rp otency
Pritc(h15a96rc)do nsidaesdr oet dh,ge r emaatoj rityc elilnns e eodfa r icshueprpo lfty r ansudates from the
ofc ontempaourtahtrohyra st,o sttheeoh balda scth arbalco­od.
teristics almotshto osifetd heegne tniecraall T hweci htohn drofobrlmaas s mta troifcx o llagen and
connectifivber otbilsassutes . mucopolysawchcihcuahnr,di cederer st caoinnd itions,
Int hpea ssto,m oeb serhvaedsr uss petchtatethd e becocmaelsc iTfhiceeo dl.l faigbecrnli oltssote h cee ll
osteombilgaohsrtti gfirnoatmth ee loiftn hmiean rgr owa rteh fien easntod f tteanak r ea dcioaulwr istrehe f erence
vessoertl hsa attl eabsotn feo rmawtaisoc nl oseltyot hceh ornodc(yZteel a1n59d9e)Tr.h, fe i boruitlssi de
depenodentn htpe r eseonfbc oen vee ssMeilnso. tt hairste aa kam eo rtea ngecnotuiaranslade r oefa m uch
(8198g)a vtehf ei arcsctu dreastcer oifpt thvieao snc ullaarr dgiearm etetro5, 8 ±0 u4 p5A . Thefsieb rruinls
sinusionsi edvseo rraglab nudsti ndo dte sctrhibebo en e ina ldlie rctiwohnisgc,ih v aefa si rulnyi foarpmp earance
sinuusnoti1id19ls.2K oll(i81k5eh3ra) md e ntitohneemd tot hmea treixxc,ie pntt ha er enae atrhs eu rfoaftc hee
bugta vneoa ccoouftn hte cihra racteristicasr.t iccaurlatriw lhaetgrheee tye ntdob ecopmaer allel
InM inocto'nsc tehpseti, n uissaol iadr cgaep illatroty h seu rfLaicteet.tal le(. 91 58h)a vfeo utnhda tth e
charactbeytr hileza ecodkfa dditcieolfnlrasolt m h e lacokfa nyo rientoafct oilolnfa igbeinrn in losr mal
orgiastnu pp;li inse hsoarl ta,rc gaep iwliltaash ri yn glcea rtiisslu abgset biyt ruait geahdnt g le ortiote hnet ation
cellw allaly.e r Thesec evwlaelsrscce uo lnasri dleciraneridtn iigndl eaggeen earratthi(rvFiei2t (g i)a.sa n(db )).
byA sch(o10ff92 1,29 4t)o, hbies tiaoncMdya txeism,o w Them ucopolysaccocmhpaornoiefdnc eta rtilage
(1298p)o inotuettdh c eo nsidpeortaebnltoeift a hleistemy a triisnx o wk nown styon tbhee tized intracellularly.
celflomsre tapalnapdsh iaag ocayntidoit tss hi iss chaIrtia sbc e­litehvaeitdto rigifrnoamtt ehgsel ycosgoe n,
teritshtafoitrc m tsh bea soifts h ree ticulo-enadboutnhdieantln hiteac slee lp lasr,t icwuhlelanar yldiyon wgan
systoefAm s ch,wo hffihcahb se emna inrleyl attoe dc alcimfaitarbCilhxeo. n drsouiltpihna tef orwmo uld
immunitoyt hadenerdf emnecceh aniTshmecs e.l lst hgee slu bstwahnicacech t asst hgel ubee twteheen
lintihnvega scwuallawlris lb ler efertroeh de raes collafigberanin ldsg ives sttota hbei lmiatItynr ix.
endothceelllfoisra,t l h eyt haoern el y ioncn oenst actth ceh apotnce arl cifirceafteirwoeinbnl eclm e a dteo the
witthh cei rcubllaotaoindnda gr ree sponsiblec hfoarnt ghtemehsa e t urnidxe rbgeofecosar lec ifioccac.tu irosn
integorfti htveye swsaellTl h.e fyo rtmr uvea sculaTrh eg reawteeasktno efts hsce h onydtrieosi c tlsa ck
endothwehliipcuhrm e vetnhbtels o fordo bmu rstionfgc ytoplparsomciecs seso ntewo i tahln iontkh er
4 Fi2g .
Fi2g .( &a ()b ) Elcetromnic roscoJpoien.ct a rtilage.
(aN)o rmaapple arantchece o lolafg en
fibrislhso wninopg r edioanmnt notrai­e
tio(nb.R) i gahntg olrei aetnitooftn h e
collfaigbetrnoit lhcsea rtislufaarcgieen
degeneraarttihvrei( t i5s,00.0 )
x
Fi3g .
Electmricorno scTohpiesc.ek tc ioonfa
chornodcystheo,w iintgs olfac cykt o­
palmsice xpanstiool nisnw ki tiht s
neighsb.o( xu 3 r,010)
Fi4g .
Elecmtrcioorsnc oTpher.ie les lturiaotns
atv aryriensgo luotfti hoceny spt olasmic
expanswihiocnchso nnecto setaecoh­
cytteo i tism meadtein eighsb aonudr
tot hen eareosstt eobl(aasT)th se.
retcrtaiooftn h oes teoaclywltsaoe c lear
vieowf t hee xpnasio(nsbA.) c yto­
palmsice xpansiisso ene bnr naching
twic(ec.D} e monrsatttehse size of the
expansAilolt nhse.se exp ansriuonn s
itnob oncea naliculae.
( 4,2;0 045,00)
x x
(Fi3g)w.,h ich expalssa ionoasnsc awlhcyi,fio cfa tion
thmea trtiaxkp elsa tchece,e lblesc owmael lienCd.u t
off frtohmte i ssuet hfcelh uoindydstrd,eoe cgenerates
anddi eSst.u doifte hsue l tra-sotftr hucech tounrder o­
cybtyee lecmtircorno (sZceoplya1 n59d9he)ar v,se h own
thasth ocrytt oplparsomciepcse sneestt rhaceta ep sule
oft hcee bluldt on octo nnweicttthh p er oceosfos tehse r
cells.
Undeorp timluomcc aoln distoimocena sr tciellalgse
mayr emacianp aobfal cet diivvei fsoiaron n s upnecified
lonpge riooftd i me. istS huceca hsw ei tthhc ee lolfs
thgee rminaolft h glera oywcetarhr tdirileacgcteol,vy e red
byac eiolfri ongbb ulsotvo eds pslealucsne ddt ehrte h ick
roofofr mebdy t htee rmibnoanplel a(tFei 1g63.) .
Thiem portoaftn hgceee r mcienlaillnts h mee chanism
ofg rowwtihbl els hoiwnnC hap1t.4e r
Thefa ctt hatc htohonecd yritsae l mosvte geat able
ceelxlp liatuisnn iscq aupea ocfbie tiyan bgtl oew ithstand
lonpge rioofdf sl usitda gonna.Ut nfiortunaantde ly,
despiititsme p orttahneicrsnee ,o e xakcntol wedogfe
thtei maec articlealgle r emmaayai lnia vfetb eeri ng
totcaulotlfffry o m niuttsr iTehndeti sffil.ct iuyds u teo
thlea cokfar elitaebclhent iodq eutete hcdete aotfth h e
celble forsei gonfsd ecompoasripetr ieosneB nyt .
histolmoegtihcoadlss ,i gmtnahsyte askese e vedraayls
toa ppear. ceUrntdfaaeviron u racbolned itions of
temperhautimudrieetc,ty. c,,a rtciellalgses umravyi ve
foar l ontgi mwei thmoeutta broelniet;we ia nsl uch
circumsast taanrtceees se mhbilbienrgnh aastsei iton n.
Foerx amtplhieen, t errouftp htbeil oofnol dot wh rough
thfee mohreaalid nP erthes'w hdiiccsahue sateshsee ,
deaotfhm osotf i toss teobolnalesyxt cse,p tionally
damatghegesr owctahr tainltdah gjeeo icnatr tiIlta ge.
is prtobhaaabtbl oe1u0 t5 h ouarrsre e qutiork eidl l
ac hornodcwyhteein tn so rmnault raireneno attvs a ilable.
OSTEEONGESIS 5
TheO steoblast Fi5g .
Anyc etlhlha atts h perp oeyro tfb eianbgl el adyto ow n Eelctromnic roscoAp es.pu erficial
osteociysst eee ant t he otfot ph e
a matwrhiicxca hl cifiweist,hd oyuitin ngt hper ociess s, picturiegn csaye ptnolsdamipcor cetsos
ano stesotbW.lh ailnoet dibffaesriicfrnaoglm tl hye ano steobrsletasiotnn gt hseru faocfe
chondrionbi ltaasbs itlt ioft oyra mc alcimfiaatbrliex , thlea mel(l 3rex,0. 00 )
idti fffreormst hacteil nil t usn iqaubei tliopt ryo duce,
orb etrteetra tihcney, t oplparsomciewcsh sielcsih n k
eacche tlol nietisg hbTohuurwssh,.e c na lcifoifc ation
thmea troicxc utrhsoe,s teobbelcaosamtneo ss teocyte Fi4g .
anrde maailnidsve es ptihtee calocfit fhiec amtaitorni x
(Fi4g).L. ei(d81y4 9n)o ttehda t the bone canaliculi
formetdht eia mtwe h en calcipfliacacanettd ih oant took
thewraens o t raocfce a nalointc huueln ic alsciidfei ed
oft hbeo ncee llTsh.ei nterceexlplaunlsraiuron n s
instihdseey stoefcm a nalwihciuiclithibs , e liaelvleodw,
thpea ssoafgn eu trienttsh ceer loelux npda nsions.
Howevietir sm, o rlei ktehlayt octchuwirisst htihne (o)
procetshseemss ienla wv aeyrs e miniosftc hetenr ta ns­
missoifno unt ribeytn htaess tropcryotcieocsf ts hees
centnrearlvs oyusst em.
Theo steoscyyntceyi tssi ouw me lplr esebrytv heed
calcifoiftc hmaeat titrohiniax rtt e stihsmeta sn ipulations
ofh istoilnocglyut,dh ifein xgoa ndt,ei calciafnidc ation
sectiootfnhsiepn egc iTmheimnsasby . e t hmea irne ason
whyt hoes teoscyyntciycit sti huoemn loyn aec cepted
bya lwlo rkeTrhse.r e is lesse xsiusptpeonrcte for the
ofc ytoplcaosnmniecc tions between osteoblasts and
osteocByutetev seh.na d tphreosceen sosbtee ser ne ­
peatseede(lnFy i ga.n pd5h )o togrtahpechyoe udnl,od t
bed enioendct eh per ocelsisnekosis ntge oocnyteto e s
anothhaedr breeecno gnizeTdh oets ot eeoixcsiy stte.
buatn o steotbrlaapsptet dh ceab lyc miaftireIidfx .
(b) (c)
Fi3g .
Fi5g .
6 thcee iltslo s urvaitav laels ,y ncymtuisoutfm necessity
bep resbefeonrtce a lcifioccactuTirhosebn. i zairdreea
Fi6g .
thtahtce y toplparsomciemcsa syde rsi ll tihnetmos elves
Photomicroofag nra arepoahfh etero­ boniets o rbjeee cted.
topicf orbmoanteii naor na bbistk 'idyn e
3m ontfholsl owliginatgi ooftn h me ain Thoes teoibnsle avsseetret nod iviTdheih.sa g si ven
renal v(e 1sx03s )e ls. ritsote h bee ltiheiafmtt u sotr igbiynt ahdteie v iosfi on
othceerlp lrse sientn htbe o nmea rrionlw a rnguem ber
Fi7g . (Weiden1r29e3i)Ic.tih s, yaeustn decideodf which
(a)P hotomicroofgw roavpebnho ne thsee vecrealollc sc uptyhimena gr rsopwa ciests h e
shwoignt hlea rcgreo,w dierrde,g ularly progietnoorft hoes teobulsaiPsnrtgi, t 'csh(9a 1r5d6)
plcaedo steoc(ytbTe)hs es. a mhesi to­
logfiiecl pdh otogrbaypp hoelda rized namet fhoecrsee l ls.
ligshhto,w liancogkf o riaetnitooftn h e Oft hopseeo pwlheoi nt hpea shta vaet tributed
fibers(. 130)
x osteogpeonteettniotcc hyv e a sculamre nttwiisioslnul e
bem adlea ter.
Thes urvtiivmoaefla no steoabflthaeasrvt i bnege n
Fig. 6 cut offi tnfrsuor tmi einspt rso balbelsys ttwhelavne
hourTsh.ee asiweasyot fk illtihen g ositseo,of c yte
courtsoiet n,e rrtuhcpeit r cuilnta htceia opni ilnli atrsy
immediate neighbotuhoers htoeoocdca.yn t Heo wever,
readbiekl ieyld al lsboy tohcecl uosfti hoen canaliculi
conneicttto it nhvgee sseelv eaInnb ,de l ibeytv hee,
breakapianorgfit t csy taospmlciocn necftriotomhn es
celilnts hm ea rrsopwa ce.
THEO STEOBLASITNWS O VENA ND
LAMELLABRO NE
Twot ypoefbs o naer gee nerdaelslcyr ibeAd,. Type
wovefinb,r oruosu,gp hr,o visbioonneB.a,l la mellar,
Havermsaitaubnro,en e.
A.W oveBno ne
Fig. 7 Thiosc cuarsas p relimsitnatagoHre ay verasnida n
lameblolnaiernt heea rclayl cifoiftc haaetn iloang e,
in etahrel y osffta racgteru,er pe aainrod,n o ccasions
inh eterootsospiifcis cuacathsim oynoss oistsiisfi cans
oro ssificoafrt eintoainls sSuaec.e radnodFt rtait tin
(1 092f)o utnhdab ty, tyirnegnv aetlsh soeen ol nsse i de
iny ounrga bbciatlsc,i fiocfta htneie ocnr toitsiscu e
occurarnetddh abto ne-wobvoenne -appaedajrae­d
centtot hcea lcdieupmo tshirtmesoe n tlhast er6 )(.F ig.
Li(e9k10 r8e)p otrhtaetwd a ahsbe tl ooe b tbaoinnfre o m
16t o2 0d ayasf tlegira titnhrege naarlt wehreytn h e
kidnheaybd e ewnr apipneo dm enttuosm e caufr reee
cicrulation.
Sintchee wsoer wkesr peu bligsrheeeaffdto rhtasv e
beemna dteo fi ndo utth nea tuorfte h set ilmutuso
ossifibcuasttof i aotrnh ey hbaeveesn u ncocutel s.s f
OsteobilnWa osvteBsno ne
Theasreoe fv arysihnagpa enssdi zaensd iarrreeg u­
larly placam eods,a sitfcro urcwmtiiutncrhgaei l nc ified
matrTihxec. o lliasgle ani d wdiotwhnao nuyptr e­
domindainrte catnifdoo nram d se ntseex t(uFri7eg) .,
whiecxhp lwahiytn hssti ypoefb oniesn amefidb rous
orr ougCha.l cifioccactuiirnaos s n c attierrreedg,u lar
manneBrl.ov oeds saerfleo su inntd h nee ighbourhood
bunto t necessariltyot hbceol noWeso.ev leyn attached
bonceo ntaaicnimsdu cpoolysaccehvaewrnhi edne s
incluidnte rda beocflu almree blolnaeTr.h ecya nnot
bed emonsitnrl aatmeebdlo lnaer eixncc eempeltni tn es
(CurarnaCdno ll1i59n7s),.