the influence of the nervous system.pdf

7/27/2019 THE INFLUENCE OF THE NERVOUS SYSTEM.pdf

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TH E INFLUENCE OF THE NERVOUS SY STEMUPON TH E GROW TH OF BONES

P . A . R ING , REDH ILL , SURREY , ENGLANDF orm er/I L am ning E van s Senio r O r thopaedic Re sea rch F ellow , Ro yal Co lleg e o f Su rg eon s of E ng land

The fo rm of th e lon g bones in the adu lt d ep ends up on th e in terac tio n of gene tic andenv ironm en ta l fac to rs. D is tu rbances of gene tic fac to rs le ad to the m ost b iza rre de fo rm ities .C h anges in the en v iro nm en t o f the bo ne usua lly p rodu ce on ly m inor chang es in the ske le ton .The im portan ce o f th e gene tic fac to r has been str ik ing ly illu stra ted by M u rray and H ux ley(192 5). B y graf ting th e p oste r io r lim b bud ofa fou r-day ch ick em b ry o on to the cho rion o f ano lder em bry o th ey show ed tha t the fem ur, and even the p ate lla , w ou ld deve lop ind ependen tlyof ex trin sic fac to rs . S im ila r g ra ftin g experim en ts by W illis (1 936 ) con firm ed the se lf-d iffe ren tia tio n of bon e. E ven in iso lated tissue cu ltu re th e knee jo in t w ill deve lop (F ell andC an ti 193 4) . In the ch ild w ith a cong en ital sp ina b if ida , even if th e lim b is anaesthe tic andcom ple tely p ara lysed , th e fo rm of the b ones is n o t g reatly d istu rbed , a lthou gh none o f th eno rm al m ech an ical s tre sse s m ay have b een ex pe rienced .

T he g row th of th e sk ele ton is p ro fo und ly in f luenced by chang es in the circu la tingho rm ones, an d by v itam in de fic ienc ie s . In con tra st w ith these facto rs the de fec ts p rod ucedby abnorm alitie s o f func tion and d istu rban ces of innerva tion appear to be sligh t. W ith no rm aln u tr itio n and en doc rine en v ironm en t, ske le tal g row th is large ly ind ep en den t o f ex te rn alfacto rs. C erta in phy sica l ch an ges in ch ild ren and adu lts m ay pro duce m od ifica tions o f sh ap e.T he ea rly w ork of v on M eyer (186 7) an d W olff (1 869 ) o n the a rch itec tu re o f the up pe r endo f th e fem u r show ed how its struc tu re w as c lose ly re la ted to m echan ica l s tre sse s. F u rth erm a them a tica l ana ly ses o fthe fem ur by K och (19 17) led to th e gene ra lisa tion s w h ich he te rm edth e law s of bon e a rch itec tu re . A t tha t tim e the ch an ges in sh ap e due to d isease w ere o n lybrie fly cons ide red .

T he d is tu rb an ces o f bon e grow th assoc iated w ith ab norm a litie s o f lim b inn erva tion areprob ab ly the comm onest cause of inequa lity o f lim b leng th . In sp ite o f th is , little is k now nabou t the m ann er in w h ich th e ne rv ous sy stem in fluences g row th . Th is is a f ield w hich len dsi tself to experim en tal and c lin ica l ex am in ation . S om e n erve les ions w hich a re assoc ia ted w ithd istu rb an ces o f b one g row th in th e ch ild can be sim ula ted in the ex pe rim en ta l an im a l. W orkin b o th th ese fie ld s has ind ica ted w ays in w hich the nervou s sy stem m ay in fluence the s tru c tu reand g row th o f bo ne .

LES ION S O F NERVE AND M USCLE iN THE EXPERIM ENTAL ANIM ALThe h in d lim b of the rabb it is a co nven ien t site fo r the p rodu ctio n of exp erim en ta l ne rve

lesio ns. T he comm on p eron ea l (sc ia tic ) n erve can be ex p lo red eas ily in the th ig h , and thepe ro neal ne rves, w h ich co n ta in m oto r fib re s fo r th e m u sc les o f th e leg and foo t, can besepa rated from th e prin cip a l senso ry ne rv e, th e su ra l. A lth ough reg en e ra tion m ay still o ccu r,even if cons ide rab le leng th s o f n erve are re sec ted , tran sm iss ion o f im pu lses d oes no t re tu rn ina y oung an im al befo re g row th stop s. T he effec ts o f m uscle inac tiv ity in the g row ing an im a lcan also be as se ssed in the young rabb it by d iv ision of th e tendons a ro und the ank le. W hena su i tab le len g th o f tend on is resec ted fu nc tion al recon stru ctio n does no t occu r.

T he rabb it is u nsu itab le fo r th e ana lysis o f the e ffec ts o f le sion s o f sp in a l ne rve roo ts ,because ofthe sm all s ize o fthe nerves invo lv ed , and becau se o fthe unusua l m ode o f d is tribu tiono f th e pe riphe ral v iscera l ne rv e fib re s in the an im a l. In the k itten , how eve r, it is p ossib le top rod uce n e rve ro o t le sio ns w ith ou t affec ting th e au to nom ic ou tflow . H ow ev er , a s w ill be show n ,bo ne m atu ra tion in th is an im a l occu rs ear ly .VOL . 43B , NO . I, FEBRUARY 1961 121


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122 r. A . R INGT he d istr ibu tio n of the m oto r , senso ry and au ton om ic ne rve fib re s to the h ind lim b of the

d og is w e ll know n , an d se lec tive les ions o f ne rv e roo ts , o r o f the sym pa the tic cha in , can b eprod uced easily in the pu ppy . E xp erim en ts in th is an im a l hav e been u sed to co nfirm tho seup on th e sm a lle r spec ie s , an d to de te rm ine m ore c lea r ly the in flu en ce w h ich d ifferen t p arts o fth e n ervo us system ex ert u pon the g row th of b one .

METHODIn each an im a l o ne s ide w as sub jec ted to op e ra tion an d the g row th o f the tib ia o n th is

s ide w as record ed , the o ther lim b se rv ing as a con tro l. S ince the tib ia is a rela tive ly straig h tbo ne , changes in g row th rate a re easily reco rded . M easu rem en ts o f tib ia l len g th a t the sta r to f the exp erim en t w ere m ade from la tera l rad iog raphs at a d istan ce o f s ix feet. In th e rabb itthe se m easu rem en ts w ere con firm ed to be accu ra te to #{216} 5 illim e tre. In th e p uppy , becauseo f the irregu lar ity o f the ep ip hy sial line , th e accu racy w as less . The fina l leng ths o f the bo nesw ere reco rd ed by m easu rem en ts w ith ca lipe rs upo n th e d ried sp ecim en s, w h ich are m ore

TAB LE IEFFECT S O F EXPLORAT ION OF COMM ON PERO NEAL NERVE U PO N LENGTH OF

TH E TIBIA IN THE RA BB IT (SHAM OPERA T ION )

D ura tion F ina l leng th in m ill im etre s D if fer ence in m illim etresumbe r (weeks) Exp erim en ta l C on tro l

72 8 2 44 .5 4 4 .5 n il73 2 2 34 .0 34 0 n il733 2 38 0 37 .5 05730 3 47 .5 47 .5 ni l73 1 3 40 .5 40 0 0. 5782 4 690 690 n il783 4 710 710 n il72 3 5 610 6l0 n il72 4 5 675 67 0 0. 5727 5 49 .5 4 9 .5 ni l72 5 6 685 68 0 05726 6 635 63 O # {216 }576 2 6 730 73 .0 ni l735 7 66 5 665 n il73 6 7 765 7& 5 n il7 63 8 840 840 n il76 4 8 725 725 ni l761 16 1020 1030 -10

Tota ls . . l,l290 l , l27 5 15

accu ra te than m easu rem en ts from rad iog raphs. F rom th ese tw o m easu rem en ts the am oun t o fg row th o ccurrin g in each tib ia w as ca lcu la ted . In th e Tab le s th e d iffe rence in the g row th rateon the tw o sid es is re ferred to as the ga in in len g th o f the a ffec ted lim b , and is expre ssed as apositiv e o r a nega tive qu an tity .

I t m u st b e em phasised th at a ll m easu rem en ts re fe r to the sha fts o f th e d ried bo nes, anddo no t inc lud e the b ony ep iph ysis . T h e ex pe rim en ts the re fo re rela te on ly to d iaphy sia l leng th s,and do no t n ecessa rily re flec t chang es in to ta l bon e d im en sion s. S im ilar ly the w e igh ts o f thedried bo nes, w hich w ere recorded afte r rem ov a l o f the fa t, and d ry ing a t room tem pera tu refo r seve ra l m on ths, refe r on ly to the d iap hysis .

S ince m any o fth e ch an ges a fter ex pe rim en tal lim b dene rv atio n m ay be re la ted to inac tiv itythe an im a ls w ere pe rm itted to run free ly du rin g the ex pe rim en t. Y oung rabb its w ere k ep t insm a ll g roup s in la rg e cages; k itten s and pupp ies w ere a llow ed th e freedom o f a la rge room .D ifficu ltie s from cro ssin fec tion w ere consid e rab le ; an im a ls dy in g p rem a tu re ly , and thosew ith trop h ic u lcera tion , have been ex clu ded from cons ide ra tion .

THE JOURNAL OF BONE AND JO INT SURGERY


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TH E. IN FLUENC E OF THE NERVOUS SY STEM UPON THE GROW TH OF BONES I 23TABLE II

EFFECT OF EX PLORAT ION OF COMM ON PERONEA L NERVE U PON DRIED W E IGHTOF TH E T IB IA IN TH E RA BBIT

N be Dura tion W e igh t in gramm es D iffer ence in gramm esm r (wee k s ) Experim en ta l C on tro l (C on tro l-E xp er im en ta l)

728 2 043 043 n il7 32 2 024 025 001733 2 032 032 n il730 3 056 057 00 1731 3 031 032 00178 2 4 232 225 -0 07783 4 234 23 7 00 372 3 5 11 9 122 00372 4 5 140 141 00172 7 5 085 086 001725 6 164 156 -0 08726 6 131 127 -00 4762 6 246 252 006735 7 175 175 n il73 6 7 158 157 . -00 176 3 8 3#{149 }4 5 3 48 003764 8 237 240 003761 16 5#{149}35 516 -019

To ta ls . . . 2987 2971 -0 16

TABLE IllEFFECT OF SUR AL NERV E D IV IS IO N U PON G ROW TH OF THE TIB IA IN TH E RABB IT

D ura tion In itia lleng th in m ill im etre s F in al leng th in m illim etr es G row th in m illim etr es G ain ofNumber a ffe cted lim b(weeks) Experim en ta l C on tro l E xp er im en ta l C on tro l Experim en tal C on tro l in m illim e tres611 2 635 635 715 715 80 80 n il63 0 2 40 0 40 0 50 0 50 0 10 0 100 n il701 2 480 480 515 515 35 35 ni l70 2 2 480 480 525 525 45 45 ni l62 2 3 39 .0 3 9 0 51 5 510 125 120 05698 3 55.5 55 .5 59 .5 59.5 4 .5 4 .5 ni l51 7 3 410 410 645 640 235 230 0561 5 4 55 .0 550 72 0 715 170 165 0562 8 4 485 485 630 63 0 14 5 145 n il695 4 550 550 650 650 10 0 100 n il695 4 560 560 630 630 70 70 n il69 9 4 49#{149}0 49 .0 55 0 550 60 60 n il709 4 650 650 760 760 1 1 0 1 1 0 n il710 4 650 65 0 760 760 1 10 1 10 n il711 4 660 660 765 765 105 105 n il612 5 605 605 800 800 195 195 n il61 6 5 630 630 845 840 215 210 0571 2 5 610 610 750 750 140 140 n il623 6 375 37 .5 645 640 270 265 05690 6 510 510 735 725 225 215 10707 6 64 0 6 4# {14 9 }0 815 825 175 185 -1 0708 6 680 680 845 850 165 170 -0 5689 7 500 500 730 725 230 225 05511 8 550 550 910 910 360 360 n il69 1 9 590 59 .0 730 730 14 0 14 0 ni l69 2 9 515 515 910 915 395 400 -0 5617 16 - 60 5 6 05 1025 1020 420 415 05624 23 39 5 3 9 .5 10 40 1050 645 65 5 --- 10625 16 400 400 900 890 500 490 10644 16 - 59.5 59.5 1030 1030 435 435 n il646 16 . 590 590 1080 1080 490 490 n ilTotals . . . . . . . . . . . 6535 6510 25

(Us ing th e d if fe rences in T ab le 1 a s co n tro l, a nd com par ing w ith them the g a in o f le ng th in th e p a ra lys ed lim b .t- 096 and P=04 .)

VOL . 43 B , NO . 1. FEBRUARY 1961


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FidT he tib iae a f ter m oto r dene rva tio n o f the h ind le gin th e rabb it. T he re is con side rab le leng then ing of

th e bon e of th e pa ra ly sed lim b.

I 24 P. A. RING

THE JOURNAL OF BONE AND JO INT SURGERY

NERVE LES IONS IN THE RABB ITUnder anaesthesia the le f t comm on

peron ea l n erve o f n ine ty rabb its aged s ix toe igh t w eeks w as ex posed in th e th ig h an d on eof the fo llow in g p ro cedu re s w as und ertaken :I ) The nerv e w as f reed a long tw o cen tim etresof its leng th , ca re be ing taken to avo idd am ag e. Th e w ound w as th en c losed w ithou tn erve re sec tion (sham opera tio n ) ; 2 ) the su ralnerv e w as f reed from the m ain trunk , and aleng th o fo ne cen tim e tre w as resec ted (senso ryd en erva tion ) ; 3 ) th e p eron ea l ne rv es w eresepara ted from the sura l an d a len g th of on ecen tim e tre resec ted (m o to r d ene rva tio n ).

Th e an im a ls w ere k illed a t p eriods va ry -ing from tw o to s ix teen o r m ore w eek s afte rth e op e ra tion . C on firm a tion o f th e e ffec ts o fth e ne rv e le sion w as by clin ica l ex am ina tiond uring the expe rim en ta l pe rio d , an d g ro ssin sp ec tion o f the affec ted n erves and m usc lesa t necropsy . N o h isto log ica l p repara tionswere made.Sham opera t ion-There w ere e igh teen an im alsin th is g roup . N o d istu rbance of func tionfo llow ed the ope ratio n , and th ere w ere n om acro scop ic chang es in m usc le o r ne rve a tnec rop sy . The e ffec ts u pon the g row th of the

tib ia a re recorded in T ab le I , and the? 1 dried w eig h ts o f th e b on es in T ab le II .I . T he chang es in leng th an d w e igh t w h ichI E fo llow sim ple expo sure and free in g ofI th e comm on p eron ea l ne rv e have b een

I u sed as co n tro ls to as ses s th e sign ifi-cance of the changes w hich fo llow theo the r p roc ed ures .

S en so ry d en erv atio n-T her e w ere th ir ty -on ean im als in th is g ro up . T he re w as no d istu rb -

ance of func tion o f th e affec ted lim b , and apart f rom an occas iona l an im al w ith paw ulce ra tionexc luded from the se rie s , the re w ere n o m acro sco p ic chang es a t n ec ropsy . T he va ria tionsin len g th be tw een the tw o tib iae a re recorded in T ab le III . B y com pariso n w ith the d iffe rencesreco rd ed in the con tro l g ro up (T ab le I) it c an be show n tha t re sectio n of the su ra l n e rvep rod uces n o sign ifican t chang es in len g th (t=09 6 , P=04). Th e w e igh ts o f the tib iae a rereco rd ed in T ab le IV . B y com pariso n w ith the d ifferences reco rd ed in th e co n tro l g ro up(Tab le II ) it can be shown tha t no sign ifican t change in w eigh t has occu rred (t=028 , P 08) .M oto r d en erv atio n-T here w ere fo rty -one an im als in th is g rou p . In these an im als a ll them usc le s a ris in g b elow th e kn ee w ere pa raly sed , and ac tive con tro l o f the foo t w as lost. A s are su lt the a ffec ted lim b w as dragged beh ind th e body . A t n ec ropsy th ese m u scle s w ere fou ndas th in fib ro -fa tty cords , o ften produc ing con siderab le defo rm ity of th e foo t.

T he len g ths o f th e tib iae a re reco rd ed in Tab le V . In th irty - th ree an im a ls th e tib ia o fthe d en erva ted lim b w as lo nge r than its fe llow , in tw o it w as sho rte r, an d in the o th e r sixthe bo nes w ere equ al in leng th . A ltho ugh the d ifferences app ear to be sm a ll, they a re


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THE IN FLUENCE O F THE NERVOUS SYSTEM UPON THE GROWTH OF BONES 12 5imm ed iate ly appa ren t w hen the d ried bo nes a re exam in ed (F ig . 1 ). W hen these d iffe rencesare com pared w ith tho se reco rded in T ab le I , the ga in in leng th o f the pa ra lysed lim b can beshow n to b e sign ifican t (t 4 .68 , P


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12 6 P . A . R INGTABLE V

EFFECT OF DIvIs IoN OF BOTH PERONEA L N ERV ES UPO N GROW TH OF THE T IB IA IN TH E RA BBIT

Dura tion In itia llen gth in m illim e tre s F ina l leng th in m illim etr es G row th in m illim e tres G a in ofNwnbe r k affec ted lim b(w ee s ) Experim en tal C on tro l Exp er im enta l C ontrol Experim en tal C on tro l in m illim e tres505 2 59 .5 5 9 .5 690 680 9#{14 9 }5 85 10507 2 59.5 59 .5 725 71 5 1 30 120 10516 2 410 41 0 53 .5 53 .0 125 120 05663 2 315 315 460 455 145 140 0570 4 2 485 485 54 0 535 55 50 0.5513 3 43 .0 430 65 5 645 225 215 1051 4 3 390 390 620 615 230 225 05510 4 570 570 740 730 170 160 10515 4 425 425 565 555 140 130 1060 9 4 540 54 .0 695 685 15 5 145 10643 4 59.5 59 .5 790 780 195 185 1065 9 4 660 660 815 810 155 150 0566 4 4 305 305 540 525 235 220 15683 4 560 560 635 615 75 5. 5 2068 4 4 56 0 5 60 685 674 125 1 15 10685 4 57 .0 570 745 730 175 160 15688 4 54.5 54 .5 66 5 65 5 120 110 10703 4 47 .0 4 70 57 .0 58 0 100 1 10 - 1050 9 5 60 0 6 0 0 835 825 235 225 1051 2 5 57 .0 570 805 780 235 210 2564 5 5 620 620 815 800 195 180 15670 6 565 56 5 790 785 225 220 05503 6 * * 865 845 * 20605 6 600 7 7 .5 7 7 .5 17 5 175 n il6 42 6 5 80 580 735 720 15 5 150 05667 6 32 5 325 805 795 480 470 1 0668 7 320 320 65 5 6 50 335 330 05669 7 320 320 665 635 345 315 30682 7 690 69 0 895 890 205 2 00 05506 8 560 560 72 5 725 165 165 n il627 8 505 50 .5 7 40 725 235 220 15756 8 57 .0 57# {149 }0 840 830 270 260 I 060 2 9 68 5 685 920 920 235 23 5 ni l7 57 9 6 0 5 6 0 5 815 800 210 195 15666 11 300 300 75 0 73 .0 45 0 430 20665 12 340 34 .0 82 0 79 5 48 0 45 -5 25758 16 650 650 95 0 95 0 300 300 n il7 59 16 635 635 9 (J .5 89 5 2 70 260 10760 16 600 600 925 925 32 5 32 5 ni l629 16+ 39 0 39 .0 9 7 .5 985 585 595 -1 0603 52 740 740 1 140 1 140 40 0 400 n ilTota ls (e xc lu d ing 503) . . . . . . . . 9160 8810 350

* Indicates no m easurem en t ava ilab le . O ve rg row th =4 per cen t.(Us ing th e d iffe re nce s in T ab le I a s con tro l, a nd com pa rin g w ith them th e ga in in len g th o f the pa ra lyse d lim b ,

t=468 a nd P =


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THE iNFLUENCE OF THE NERVOUS SYSTEM UPON THE GROW TH OF BONES 12 7For th is se r ies o f an im als no d irec t con tro ls a re av ailab le , an d to consid e r the va lid ity o f theobserva tion s, m easu rem en ts from the con tro l g ro up of rab b its have been used . W ith th ism e thod , th e g ain in len g th o fth e p a ra lysed lim b has no statis tica l s ign if ican ce (t= l21 , P=02).

The w e igh ts o f the b ones in th ese an im a ls are reco rded in T ab le X . Th e bone o f thep a ra lysed lim b w as lig h te r than its fe llow in th ir teen . U sing th e m easu rem en ts o f bo ne w e igh t

TABLE V IE FFE CT O F D IV IsIoN O F BOTH PERON EAL NERV E S U PON W E IGHT OF THE DR IED

T IB IA E IN TIlE R ABB IT

N be D ura tio n W eigh t in gramm es D ifference in g r ammesm r (wee k s ) Exp er im enta l C ontrol (C on tro l- E xp er im enta l)

50 5 2 148 163 01550 7 2 174 186 01251 6 2 086 113 027663 2 065 068 00370 4 2 065 068 00351 3 3 145 l67 02251 4 3 12 7 150 023510 4 18 2 23 0 04 851 5 4 103 119 01660 9 4 163 167 00464 3 4 244 294 050659 4 224 241 01766 4 4 080 085 005683 4 086 088 002684 4 15 5 158 00368 5 4 17 1 20 6 03 5688 4 156 173 01770 3 4 079 084 00550 9 5 247 294 04751 2 5 26 4 287 02364 5 5 279 3 0 5 02 6670 6 215 232 01 7503 6 225 25 5 03 0605 6 250 237 - 01364 2 6 180 200 020667 6 115 125 010668 7 090 112 022669 7 100 106 006682 7 225 253 028506 8 160 157 -00 36 27 8 161 169 008756 8 212 243 031602 9 331 350 01975 7 9 18 1 23 4 05 3666 II 195 216 021665 12 157 165 00 8758 16 3 3 3 3 .9 5 06 275 9 16 255 290 03576 0 16 238 307 069629 16+ 322 397 0 7560 3 52 563 632 069

Totals . . . 7751 8721 9 .70

(Us ing th e d iffe re nce s re co rd ed in T ab le II a s a con tro l, t= 6 3 and P =


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12 8 i . A . R INGTABLE V II

EFFECT O F TENDON DIV ISIO N UPON GROW TH OF TH E T IB IA IN THE RA BB IT

Du ra tion ln itia lleng th in m illim etre s F ina llen g th in m illim e tre s G row th in m illim e tre s G a in o fNumbe r k af fected l imbwe e 5, Exp er im en ta l C on tro l Experim en tal C on tro l Exper im ental C ontrol in m illim e tres647 2 505 505 585 570 80 65 1565 0 2 51 5 51 5 # {2 16 } 59 .5 85 8 0 05754 2 57# {149}5 57 5 610 610 3 .5 3 .5 n il75 5 2 55#{149}5 55 .5 57 .5 57 .5 20 20 n il78 1 3 570 57 0 695 690 125 l20 0566 2 4 53 .5 5 3 .5 76 5 74# {14 9 }5 23 O 210 20751 4 490 490 615 595 125 105 20753 6 520 520 755 74 .5 235 225 1077 8 6 * 79.5 78 5 * * 1077 9 8 480 480 830 820 350 34 -0 1 0780 8 * 820 790 * 30651 9 * * 87 5 85 5 * 20672 12 570 55 5 920 880 35 0 325 25677 12 550 55 0 910 895 360 345 15678 12 535 53 .5 79 .5 7 65 260 230 30675 16 650 65 0 91 5 91 0 2 65 260 05676 16 560 560 985 975 425 41 5 1 0

To tals (ex clu d ing 7 78 , 7 80 , 65 1) . . . . . . . 2945 2775 17 0* Indicate s no m easurem en t a va ila b le .

(U sing the d iffe rences in Ta b le I a s a con tro l, a nd com pa rin g w ith them th e ga in in le n g th of t he p ar al ys edl imb , t=2 8 , P=


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1: JoF IG . 2

Th e tib iae a fte r a n te rio r ro o t les ion in th epupp y show in g d im in u tion in sha ft d iam e te r,a nd d im in ut io n in to ta l le ng th b ut an i nc r easein d iaph ys ia l leng th on the s ide o f th e les ion .

THE IN FLUENCE OF THE NERVOUS SY STEM UPON THE GROW TH OF BONES I 29

VOL . 43 B , NO . I, FEBRUARY 1961

JoFIG . 3

T he t ib ia e a fte r a n te rio r n e rv e roo t le s io n inth e puppy . The re is o ve ra l l le ng the n in g o fthe bo ne on the p ara lys ed s id e a nd a m a rke dd i f fe rence be tween the s ize s o f th e bon y

ep iphy ses .

d is tu rbance, and in the am oun t o f m u scle w as ting . In tw o an im a ls pa ra lys is ap peared to becon fin ed to the toes, w he reas at the o th er ex trem e th ree an im a ls exh ib ited a flacc id p ara lysiso f all the h in d lim b m usc le s .

The re w ere tw en ty -o ne an im a ls in th is se rie s , and each w as k illed th ree m on th s a fte rthe ope ratio n . Th e d ried bo nes show ed w astin g w h ich w as in gene ral p ropo rtion al to theex ten t o f the m u sc le pa raly sis . M easu rem en ts o f bo ne w eig h t w ere n o t m ade , since in m anysp ec im ens th e ep iph ysis rem ain ed a ttached to th e sha ft o f the b one . M easu rem en ts o fd iaph ysia lleng th a re p resen ted in T ab le X l. F if teen an im als show ed p ara ly tic bo ne leng then ing , an d inthree o th ers the bon es w ere equ a l in leng th . T here be ing no m o re su itab le co n tro l ava ilab leth e sign ificance o f th ese observa tion s h as been tested aga inst the ch an ges in len g th w h ichfo llow sym pa thec tom y in the puppy (see T ab le X II). U sing th is m e tho d t2 4 , P


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N um ber S urv iva l(weeks )Initiallength in m illim etres F ina llen g th in m illim e tre s G row th in m illim etr es

Exp er im en ta l C on tro l E xperim en ta l C on tro l E xp er im en ta l C on tro l

Leng then ingo faffected l imbi n m i ll im e tr e s

837 3 6478 9 4830 4 60 583 2 4 63080 5 5 61 081 0 580 2 6 70#{149 }0803 6 62080 4 6 675806 6 750808 6 570809 6 55 0811 681 2 6813 6 73#{149}0814 6 660

64 0*

60 563 061

*

70 062 067 575 057#{149 }055 0

**

73 .066 0

65 542 561 564 565 55& 073 .56& 571 581 064 563 062 068 075 .571 0

65 542 561 564 565558 073 .568 070#{149}581#{149}063 562 562 067 075 070 5

15 1 .5 ni l* * ni l10 10 nil15 15 ni l4#{149}0 4 .0 n il* * ni l3 .5 3 .5 n il6 5 60 0540 30 1060 60 n il7.5 65 1080 75 0 .5* * ni l* * 1025 20 055#{149}0 4 #{1 49 }5 0# {14 9}5

To ta ls (e xc lud ing 7 89 , 8 10 , 8 11 , 8 12 ) . . . . . . 510 470 40

D ura tion W eig h t in gramm es D ifference in gramm esNumbe r (weeks ) - (Control-Experimental)Exper imen ta l Cont ro l837 3 134 145 011789 4 036 040 00483 0 4 157 145 -0 12832 4 15 5 158 003805 5 167 176 00981 0 5 096 095 -0 01802 6 215 245 030803 6 145 l6 0 015804 6 177 206 029806 6 221 255 034808 6 116 135 019809 6 081 124 04381 1 6 14 2 13 2 -010812 6 157 195 038813 6 228 234 006814 6 174 186 012

Tota ls . . . 2401 2631 230

I 30 P . A . R ING

THE JOURNAL OF BONE AND JO IN T SURGERY

TA BLE IXGROWTH O F T iB IA A FTER AN TER IO R LUM BAR ROO T D IV IS ION IN K Ir rEN s W E IGH IN G I- I 5 K ILOGRAM S A T OPERA T ION

* Ind ica tes no m easu rem en t a va ilab le .(Us ing the d iffe ren ce s in Ta b le I as co n tro l, and com par ing w ith th em the le ng the n in g o f th e p a ra lys ed le g ,t= l21 and P =02 .)

TABLE XW EIGH T O F DR IED TIB IAE A FTER ANTER IOR LUM BAR R ooT D IV IS IO N IN K I-rrEN s

WEIGH ING 1-15 KILOGRAMS

Us in g th e d iffe re nces rec o rd ed in T ab le II a s con tro l, t= 2# {1 49 }9 3 ,P =


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THE IN FLUENCE O F THE NERVOUS SYSTEM UPON THE GROW TH OF BONES 13 1TABLE X l

EFFEC T O F A NTER IOR RooT D IV IS ION U PON GROW TH OF THE TIB IA IN THE PU PPY

In itia l leng th in m illim e tres F in al length in m illim e tre s G row th in m ill im e tre s G a in ofN um ber a ffec ted lim bExperim en tal C on tro l E xperim en ta l C on tro l E xperim en ta l C on tro l i n m i ll im e tr e s

23 82 82 142 143 60 61 -124 68 68 100 98 32 30 225 70 70 120 118 50 48 227 70 70 104 102 34 32 228 72 72 105 105 33 33 nil31 61 61 99 95 38 34 432 83 83 100 97 17 14 333 75 75 102 103 27 28 -134 65 65 109 105 44 40 435 75 75 114 111 39 36 336 64 64 97 93 33 29 465 62 62 121 121 59 59 nil66 64 64 114 110 50 46 467 72 72 125 124 53 52 168 81 81 132 131 51 50 171 80 80 106 105 26 25 172 71 71 82 82 II II nil73 75 75 113 Ill 38 36 274 58 58 94 91 36 33 375 60 60 90 92 30 32 -276 70 70 105 103 35 33 2

To ta ls . . . . . . . . . . 796 762 34

( By c om p ar is on w ith Ta b le X II , t 24 a nd P (00l.)

TABLE X IIEFFECT OF LUM BAR SYM PA THECTOM Y UPON GROW TH OF THE T IB IA IN THE PUPPY

In itia l len gth in m illim etr es F in al len gth in m illim etres G row th in m illim etres G a in ofNumber affected limb

E xper im ental C on tro l Exper im ental C on tro l Exper im ental C on tro l in m illim etres42 81 81 123 123 42 42 nil45 116 116 138 138 22 22 nil46 134 134 188 188 54 54 nil54 74 74 144 143 70 69 155 68 68 118 117 50 49 157 84 84 127 126 43 42 158 84 84 130 129 46 46 nil59 72 72 121 121 49 49 n il

632 63 63 126 126 63 63 nil633 57 57 84 84 27 27 nil634 56 56 96 96 40 40 nil

To ta ls . . . . . . . . . . 50 6 50 3 3

There w ere eleven an im a ls in th is g roup and the g row th of th e lim b s is reco rd ed inT ab le X II . In th ree an im a ls the tib ia o f the sym pa thec tom ised lim b w as lo nge r th an its fellow ,in the o ther e igh t the bo nes w ere o f equa l leng th . N o sa tis fac to ry co n tro l is ava ilab le fo r th isgrou p , bu t sim p le insp ec tion o f the f igu re s sh ow s th at sym pa thec tom y is w ithou t sig n ifican te f fec t upon the g row th o f the tib ia in th e p uppy .

LOW ER MOTOR NEURONE LESION S IN CH ILDREND istu rbances o f lim b grow th in ch ild hood a re comm only assoc iated w ith ab norm a litie s

ofth e n ervous sys tem . T he defec t m ay lie in the b ra in , p rod uc ing upper m oto r neurone defec ts,or in th e sp ina l co rd , o r pe riphe ral n e rves, g iv ing rise m a in ly to low er m oto r neu rone d iso rde rs .vO l . . 43 B , NO . I, F EBR UARY 1961


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13 2 i. A . R INGA lth ough th e type of ne rv e les ion m ay va ry con side rab ly , the fina l effec ts up on th e ske leto nrem a in the sam e-n am e ly , a trop hy o f co rtica l bo ne , re ta rd a tio n of g row th in leng th , an d ach an ge in ov era ll p ropo rtio ns and fo rm . Th e sim ilar ity o f these changes in w ide ly d iffer ingd iseases sugg es ts th at the re is som e comm on facto r w hich in fluences the shape and stru ctu reofy oung bone . In o rtho paed ic p rac tice ab norm a litie s o fg row th in len g th becom e of im po rtanceo n ly w h en they a re un ilate ral, and abnorm alitie s o f fo rm are o f sig n if icance o n ly w h en th eyproduce defo rm ity . M any o f th e ef fects o f nerve lesion s a re m ask ed by the b ila tera l symm etryof the d istu rbance .

T he ch an ges in th e g row th of bon e w hich accom pany pa ra ly tic po liom yelitis a ffo rdva luab le ev idence o f the w ay in w hich the nervo us sys tem inf lu en ces the ske le ton . T hein trodu c tion of th e scano grap h has p erm itted accu ra te rad iog raph ica l m easu rem en ts o f lim bgrow th to b e m ade , and th ere is now substan tia l ag reem en t u pon the late effec ts o f m usclep ara lysis . D eta iled stu d ie s have been reco rd ed e lsew here (R ing 1957, 1 958; R ing an d W ard1958 ; R atl if f 1959 ) and on ly th e m ain fea tu res o f th ese stud ies w ill be reca lled .

The imm ed iate e ffec t o f th e neurona l le sion in po liom ye litis is to p rodu ce leng then ing o fth e bon es o f the a ffected lim b . Th is in c rea se is appa ren t du ring the firs t year a fte r the o nse to f the d isease , and m ay pe rs is t d u rin g th e second yea r, b u t the rea fte r the g row th o f thep ara lysed lim b is reduced . The d istu rb an ce of g row th pe rsis ts th rou ghou t ch ild hood , andth e d isc rep an cy in lim b leng th in crease s stead ily . If th e pa ra ly sis is m a in ly u n ila te ra l, thed iffe rence in leng th b etw een th e legs is , in gene ral, re la ted to the rela tive lo ss o f m usc le b u lk .T o th is ru le the re are occasiona l excep tion s, and th e ex ten t o f the m u sc le lo ss can no t be usedto p red ict th e d eg ree o f sh o rten in g w ith any ce rta in ty . In p ara ly tic po liom ye litis the o ccasion alp atien t w ith seve re un ilate ral p ara lysis w ith ou t sho rten ing , o r sligh t pa re sis w ith seve reshorten ing , is o ften cited as ev iden ce ag ain st the causal as soc ia tion of pa res is and re tarda tiono f lim b g row th . T h is a ssoc iatio n has w iths tood s tatis tic a l an aly sis on seve ra l occasio ns(S tinch fie ld , R eid y an d B arr 1 949 ; G u llickson , O lson and K o ttke 1950), bu t p ro bab ly th em usc le los s is n o t the o n ly fac to r w h ich de te rm ines lim b shorten ing .

In p o liom ye litis the g row th of the tib ia is usua lly a ffected m ore than th e g row th of thefem u r, rega rd le ss o f th e d istr ibu tion o f the lesio n . Indeed th ere is n o ev iden ce th at the latte ris o f g rea t im po rtance . I t is the ex ten t o f the m uscle lo ss , ra the r than its ex ac t d is tr ibu tion ,w h ich de te rm in es the fa te o f th e lim b . T he sam e is u ndoub ted ly tru e in m ye lo -dysp la sia.A lthou gh th e d istu rb an ce in these ch ild ren is a con gen ita l o ne , and the ph ase o f pa ra ly ticbon e leng th en ing is th e re fo re n o t seen , th e g row th ch anges in the lim b fo llow clo se ly th ose inpo liom yelitis . S im ila r chang es occur in the ch ild w ith a p e riph era l n e rve le sion . P erhaps thecom m onest o f the se in o rtho paed ic p rac tice is the b irth in ju ry o f the b rach ial p lexu s. W henrecov ery is rap id an d com p le te the re is little o r no shorten ing . P ersis ten t m usc le w eak nessleads to sh orten ing of the a rm and re tarda tion o f th e g row th of the hand ; the inequa lity isusu a lly o f little c lin ical s ig n ificance .

In any stud y o f the effec ts o f p o liom ye litis upo n th e g row th of the lim b a tten tion m ustbe pa id to the c ircu la to ry chang es w hich occur . G ask and R oss (19 34) a sc rib ed th ese to thelack o f s tim u li assoc iated w ith no rm al activ ity . C onsequ en tly the lum en o f the m a in v es selso f the lim b m ay b ecom e d im in ish ed (T elfo rd and S top fo rd 19 33). Th ere can be little d oub ttha t th e vascu la r req u irem en ts o f the pa raly sed lim b a re d im in ished , bu t d irec t ev idence islack ing : m o st o f the inv estiga tions o f lim b b loo d flow h ave been conce rn ed w ith the po ssib ilityo f a d istu rbance o f the au tonom ic ne rv ous system in the la ter stag es o f p o liom ye litis , and dea lin gene ra l w ith sk in b lo od flow . M cPhe rson and K essel ( 1955) inves t iga ted th e re sting b loo dflow in the fee t o fp a tien ts w ith res idua l pa raly sis . T hey found a s ign if ican t d iffe rence be tw een th eflow in th e tw o fee t w h ich they asc r ibed to a hy po th etica l v iscera l ne rv ous le sion situa ted in thehypo th alm us. If su ch a le sion do es in fac t occu r , it c an h ard ly be resp ons ib le fo r lim b sho rten in g .

G oe tz, du To it and Sw art ( 1955) inv estiga ted the c ircu la to ry chang es a fte r po liom ye litisby the op tica l d ig ita l p le sth ysm ograp h , a m e tho d w hich reco rds b lood flow in the sk in ra the r

ThE JOURNAL OF BON E AND JO IN T SURGERY


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TH E IN FLU ENCE O F THE NERVOUS SYSTEM U PON TH E GROW TH OF BON ES I 33than in m uscle . T hey sug gested tha t th ree facto rs w ere asso cia ted w ith the v ascu lar d is tu rban ces:1 ) Th e lack of stim u li p rom o ting an e ff icien t circu la tion , and d im in ished by v ir tue o f th epara lys is ; 2) in te rfe rence w ith sym pathe tic innerva tion , p robab ly caused by dam ag e topreg an g lio n ic ce lls in the late ra l ho rn ; and 3) a r ter ia l na rrow ing . The firs t and th ird o f th esefac to rs a re accep tab le , a lth ough it is like ly th at changes in the vessels are o n ly secon da ry tothe lack of n orm al stim uli rathe r than to a prim ary d istu rbance . T here is little ev id en cefrom path o log ica l s tud ies tha t p erm anen t d am ag e to the sym path etic occu rs in p o liom ye litis ,a ltho ugh th ere is som e c lin ical ev id en ce tha t sym pa the tic d istu rbances m ay o ccu r in th e earlys tages o f th e d isease (L undb aek 1943 ; Sm ith , R osen b la tt and L im auro 1949). T h e abnorm alvascu la r re spo n se s o f the late r stages cann o t be exp la ined by sim ple d estru ctio n of sym pa the ticvasocons t rict ors . In deed , the p regang lio n ic ce lls in th e la te ra l ho rn o f the sp ina l co rd a red istin ctly resis tan t to a ttack by po liom ye litis , an d on ly w hen d es truc tio n has been ex trem elyseve re -fo r exam p le , w h en th ere has been com ple te pa ra lys is o f all abd om in al and in te rco sta lm usc les -a re these ce lls a ffec ted (Sharrard 1959) . T he in fluence of the sym pa the tic sy stemupon th e g row th of bo ne w ill be d iscussed la ter , bu t it is c lear tha t its in f luence is slig h t, andin su ffic ien t to accoun t fo r the d istu rb ance of bone grow th .

The circu la to ry ch an ges in p o liom ye litis h av e a lso b een investiga ted by reco rd in g chang esin sk in an d m u sc le tem pera tu re (T ro tt, N eslin e and G reen 1958) . In pa tien ts w ith un ila te ra llim b in vo lv em en t, a sign ifican t dep res sion o f sk in an d m usc le tem peratu re s d id no t app ea run til s ix m on ths a f ter th e on se t o f th e d isea se . Th e re w as a d irec t a ssoc ia tion b etw een th ed im inu tio n in sk in and m u sc le tem pera tu re and the ex ten t o f th e pa ra lys is . I f w e assum e tha tm u sc le m etabo lism does no t co n tr ibu te sign ifican tly to the w arm th of the lim b , these ch an gesp rob ab ly in d icate a true d im inu tio n of b lood flow . S in ce it ha s a lready been show n tha tsh o rten ing and pa ra lysis a re rela ted , it m igh t be a rg ued th at the d im inu tio n of b lo od flow isitse lf re spon sib le fo r the d istu rbance o f b one grow th . T h is ex tensio n of th e a rg um en t im p lie sthat u lt im ately the b lo od su pp ly o f the ske leto n is affec ted , and fo r th is the re is a t the m om en tno d irec t p roo f.

THE RELAT IONSH IP BETW EEN THE NERVOUS SYSTEM AND THE SKELETONTHE INNERVATION O F BONE

N erve fib res can be dem onstra ted in the p erio steum , in th e bon e m arrow and in the b oneitse lf. T he pe rio steum is free ly su pp lied w ith sen so ry ne rve fib re s , te rm ina ting as unm yelin atedne rve n etw orks (F iend e l, W edde ll and S inc la ir 194 8) . M ye lina ted and unm ye lina ted fib re shave been no ted in the bon e m arrow o fy oung an im a ls by R oss i (1 932 ). F ib re s have a lso b eendesc ribed en te rin g b on e alo ng th e v ascu lar ch an ne ls (S t#{246 }h r1928), p ar tly vasom oto r, and pa rtlysensory f ib re s fo r pa in . In g row ing bone D e C astro (19 29) repo rted ne rve fib re s en d ing inclo se re la tionsh ip to the os teo b las t, bu t these fib res w ere no t fo und in m atu re bone . A ltho ughH urre ll (19 37) rep orted the p re sence of ne rve fib re s in adu lt bon e, en d ing e ithe r b lin d ly o r inrela tion to the osteocy te , h is ob se rv atio ns w ere b ased upon a sing le sp ecim en , and have n o tb ee n c on fi rm ed .

It is genera lly agreed th at no fib res a rising in the m o to r ce lls o f the sp ina l co rd te rm ina tein bon e, and the fib re s w h ich have been desc ribed are de riv ed from th e senso ry and thesym pa the tic n ervo us sys tem s. In the exp erim en ta l an im a l the in fluen ce of the m oto r, thesensory and th e au to nom ic pa rts o f th e ne rv ous sy stem can be exam in ed by se lectiv e n erved iv is ion . In m an , cer ta in se lectiv e neu ro lo g ica l le sion s a re a lso ava ilab le fo r con side ra tion .

THE IN FLU ENC E OF THE SENSO RY NERV ESCorb in and H insey (19 39) show ed tha t d iv is ion ofth e poste r io r n e rve ro o ts in the m a tu re

ca t p ro duces no ch anges in the fo rm and phys ica l p rope rties o f bo ne . To th is they m ak e oneexcep tio n : in the h ip , ch an ges m ay occur in assoc iatio n w ith a neu rop a th ic a rth rop ath y .VOL . 43 B , NO . I, FEBRUARY l96J


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13 4 P . A . R INGSenso ry ne rve d iv is ion in th e y oung k itten , a lthou gh o ften p rodu cin g co nside rab le d isab ilityand d isuse , does no t in flu ence th e g row th in leng th o r b read th o f bon es (G ille sp ie 1954) .

Sen so ry d is tu rb ances o f th e lim bs in ch ildh ood are ra re , ex cep t in a ssoc iatio n w ith am oto r d efic it. In adu lts pu re senso ry d istu rbances are m o re com m on . C hang es in the stru c tu reo f th e sha fts o f the long bones do no t o ccu r . C h an ges a t the ends o f th e bon e m ay be seen inas soc ia tio n w ith the neuropa th ic jo in t. T hese ch an ges a re p robab ly seco nda ry to the traum aas soc ia ted w ith an insen sitive jo in t, and the changes in b one s tre sse s w hich en sue . T hed iso rgan isa tion of m usc le tone and ac tio n due to pro priocep tive abn orm alitie s m us t a lso p laya part.

TH E V ISCER AL NERVOUS SYSTEMT here is little expe rim en ta l ev id en ce to sugg es t th at sym path ec tom y in fluences th e ra te

o f g row th of norm a l bo nes. C an non , N ew ton , B righ t, M enk in an d M oo re (192 9) in th ek itten , an d B acq (1930) in th e yo ung rat, fo und no ch an ges a fte r un ilate ral sym pa thec tom y inthe fina l len g th o f th e lim bs. B isga rd (1 933 ) no ted no chan ges in the g row th rate o f the tib iao f the y oung goa t a fte r u n ila tera l sym pa thectom y , a ltho ugh th e su rface tem pera tu re o f thelim b w as inc rea sed . H arr is and M cD ona ld (19 36) w ere un ab le to de tec t hy pe raem ia o f thelim b a fter sym pa thectom y in the k itten , pu ppy or lam b , an d no chang e in th e rate o f bo neg row th w as de tec ted . H ow eve r , G u llickson , K ub icek and K ottke (19 5 1 ) reported lim bsh orten in g , m a in ly in th e m e ta tarsa l reg ion , a f ter p ro lo nged stim u latio n of the lum barsym pa the tic ch a in in the pu ppy , an d lim b len g then in g (the d istr ibu tio n of w hich w as no td escribed) afte r its exc ision . G oe tz, du Toit and Sw art (1955) ca r r ied ou t a lum barsym pa thectom y in the young rab b it w h ich produ ced an e lev atio n o f sk in tem pera tu re, andan in crease o f abo u t 4 pe r cen t in the leng th o f th e p aw , th e rest o f the lim b rem a in ingunaffected.

A lth ough sym pa thectom y has com m on ly been p e rfo rm ed fo r the circu la to ry d is tu rb ancesw hich fo llow po liom ye litis , few accoun ts hav e b een pub lish ed of th e e ffects o f th is p rocedu reup on the g row th o f th e pa ra lysed lim b . H arris (19 30 ) an d O gilv ie (19 32) each repo rted apa tien t w ith reduced sho rten in g afte r lum bar g an g lion ec tom y . In 19 36 H arr is and M cD ona ldco llec ted fo r rev iew fo rty -s ix o f th e seven ty ch ild ren they had su b jec ted to sym pa thectom y .O f th ese ch ild ren 46 p er cen t sh ow ed redu ced shorten ing rang in g from one -eig h th o f an inchto o ne inch , and in 17 pe r cen t sh o rten ing fa iled to inc rea se . In the rem a in ing 37 pe r cen t thed isc repancy in len g th progressed bu t tw o-th ird s o f these w ere found no t to have a perm anen tsym pa th ec tom y . S in ce the sh orten ing in po liom ye litis is on the w ho le p ro g ress ive , 63 pe r cen to f the se pa tien ts m ay be sa id to h av e de riv ed som e bene fit from th e op era tion . In 1 950 B arr,S tinch field an d R e idy pub lished a com pariso n be tw een the p rogre ss o f lim b sh orten ing afte rpo liom ye litis in a con tro l g ro up o f p atien ts , an d in a sim ila r g rou p w ho had been su bm ittedto un ila tera l lum bar sym pa thec tom y . They fo und tha t w h e reas the inequ a lity o f lim b leng thbecam e m o re m arked in 91 pe r cen t o f the con tro ls , o n ly 57 pe r cen t o f th e ch ild ren sub jec tedto sym path ec tom y show ed prog re ssio n . In the con tro l g ro up the av e rage in c rea se in thed iscrepancy w as 1 8 cen tim e tres , in th e trea ted g ro up 03 cen tim e tre . Th us the sym pa thec tom yh ad inc reased the g row th o f the pa raly sed lim b by 1 5 cen t ime tres . Since the average age ofth e ir pa tien ts w as 83 yea rs , m any of the ch ild ren m u st h av e b een ap proach ing the end o f theg row ing p eriod and it is poss ib le th a t ear lie r ope ratio n w ou ld have inc rea sed the co rrec tionob tain ed . It m ust be rem em bered tha t the se pape rs are b ased en tire ly up on c lin ica l m easu rem en tsand in c lu de no sta tis tic al ana lysis , and tha t the da ta p re sen ted are n o t su itab le fo r ind ep en den tsta tis tic al an aly sis . M ost in vestiga to rs w ou ld ag ree th at estab lished lim b in equa lity a fte rp o liom ye litis neve r ten ds to im prove , and any m e tho d of trea tm en t w h ich d im in ishes the lim bin equa lity revea led b y the scano grap h m ig h t be rega rd ed as sign ifican t. T he re is , h ow eve r,o ne im po rtan t gap in our kn ow ledge of th e fa te o f the short lim b a t p ube rty . If the ep iph yseson th e affec ted sid e fuse la te r than no rm ally , the shorten ing w ou ld tend to reg ress at th is tim e.

TH E JO URNA L OF BON E AND JO INT SURGERY


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THE INFLUENCE OF THE NERVOUS SY STEM UPON THE GROW TH OF BONES I 35

Fu rthe r ob se rva tion s on th e na tu ra l h is to ry o f the g row th of th e p ara lysed lim b a re necessa rybe fo re o ne can be ce rta in o f th e s ign if ican ce o f the chan ges reco rd ed a fte r sym pa thec tom y .

In re latin g th e re su lts o fexp erim en ts in an im a ls to the e ffec ts o fsym pa thec tom y in ch ild renw ith m uscle p ara ly sis th e fo llow ing questions m ust b e co nside red : 1 ) Is it p oss ib le to s tim u la tethe rate o f g row th of a norm al b one , o r on ly on e which is abn orm ally sh ort? 2) W ha t is theno rm al sym path e tic inne rva tion o fthe ex trem itie s in ch ild ren and expe rim en ta l an im a ls? H owex tens ive m ust the sym pathec tom y be ? 3 ) T o w hat ex ten t is th e fun ctio n of th e au ton om icin ne rv atio n of the lim bs com parab le in young an im a ls and ch ild ren?S tim ulation o f norm al grow th -A lthough it is po ss ib le tha t the e ffec ts o f sym pathec tom y aregrea te r w h en th e g row th of a lim b is alread y d is tu rb ed , th e ob se rva tio ns o f H arris an dM cDona ld (193 6) su ggest tha t ove rg row th ofa no rm al lim b m ay occur af te r u n ila te ra l lum barsym pathec tom y fo r H irchsprun gs d isease . In each of the fou r pa tien ts ex am ined they fou ndtha t the lim b on th e s ide o f th e sym path ec tom y w as long er than its fe llow . A lthou gh th ed ifferences in th ree o f th e pa tien ts w ere co nside rab le (ha lf an in ch or m ore ) they w ere basedu pon clin ica l m easu rem en ts a lo ne . F ah ey (1 936 ) ex am ined tw o sim ila r pa tien ts b y te le -rad iog raphs and w as u nab le to de tect any d ifference in th e len g ths o f the legs .T he com parativ e innervation -T he preg an g lion ic sym pathe tic fib res innerv ating the low er lim bin the ch ild ar ise from th e low er th o rac ic and th e u ppe r tw o lum bar segm en ts o f th e sp in alco rd . A lthou gh m ost o f them descen d in the sym pa the tic trunk be fo re re lay ing in th e lum baran d sacra l sym pa the tic gang lia , som e post-gang lio n ic fib re s ar ise a s h igh as th e n in th tho racicgang lio n . R em ova l o f o n ly the secon d , th ird and fourth lum bar g an g lia is p ro bab ly anad eq ua te m e thod of p rodu cin g com ple te p regang lion ic den erva tion of the fo o t (R oss 1946).A m ore sa tisfacto ry d en e rva tion is ach ieved by ex ten d ing the o pe ra tion to th e rem ova l o f thef irs t lum b ar gang lion (M cPh erson and K esse l 19 56).

F o r m ost ex pe rim en tal an im als de ta ils o f the cen tra l o r ig in o f the sym path etic f ib re s a reno t ava ilab le . In the dog the pregang lion ic sym pathe tic ou tflow m ay con tin ue dow n as fa ras th e s ix th lum bar segm ent o f the sp ina l co rd (M ehler , F isch er and A lexander 1952 ). F o rdene rv atio n of the h ind lim b of th is an im a l rem ov al o f the la st lum bar an d the firs t sacra lgang lia is im portan t (P e rp ina 1 955 ).The com para tive physio logy -A n effec tive lum bar sym pathec tom y in a ch ild sho u ld produ cepe rmanen t in c rease in the b lood flow in sk in and m u sc les, an in crease in sk in tem pera tu re ,and absence of sw eat secre tion in the affec ted lim b . In th e fu rred an im a l the sym pa the ticsystem plays no part in the regu la tion of body tem peratu re , and the re is no sim p le m eth odof d eterm in in g wheth er an adequa te sym pathec tom y h as been perfo rm ed . T hus, thein te rp re ta t ion of an im al ex perim en ts requ ires co nside rab le cau tion . The w ork o f F e ll (194 9)up on th e k itten sugg ests th a t on e of the e ffects o f sym pa thec tom y is to p ro duce en la rgem en tof the H av ersian cana ls in bo ne . T h is canno t b e taken as pro of o f an increase in bone b lo odflow . Z in n and G riff ith (19 41) have sh ow n tha t th e ra te o f up tak e of carbo n pa rtic le s in th ebon e o f the ra t is d im in ished afte r sym path ec tom y .

F rom th e ev idence pre sen ted it is c lea r th a t sym pa thec tom y in the dog has n o sign if ican te ffect up on bone grow th . O n e canno t ded uce tha t th e sam e is true in the ch ild . A lthou ghthe ev idence is slender, and lacks rad iograph ic confirm ation , it is possib le tha t an adequa teun ila te ra l lum bar sym pa thec tom y p ro duces a sligh t inc rea se in th e ra te o f b one grow th .

THE M OTOR NERVESA ltho ugh the e ffe ren t fib re s o f the sp ina l co rd d o no t d irec tly inn erva te the ske le ton th ey

a re th e o n ly fib re s w hich exe rt any ap prec iab le in f luence upon the struc tu re and com positio nof the b one . E xpe rim en ta l w o rk upon the in f luence of m o to r n erve d iv isio n has b een d irectedm a in ly tow ards th e ch an ges in d iam e ter w h ich fo llow n erve sec tion ra the r than the ch angesin leng th . Po tto rf (191 6) and H ow e ll (1 917 ) in s ing le exp erim en ts up on the leg o f the p uppyfo und m arked a troph y and sligh t sho rten ing of lim b bon es a fte r p eriph era l ne rv e sec tion .VOL . 43 B , NO . I, FEBRUARY 1961


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13 6 P . A . R INGHow ell conc lud ed tha t th e g row th of bon e in leng th w as la rge ly ind ep enden t o f ex te rn alfactors, bu t its g row th in b read th w as c lose ly re la ted to ch an ges in stress an d s tra in .

In ad u lt d ogs the e ffec ts o f d isuse , fo llow ing n erve sec tion o r imm obilisa tion , w ere stu d iedby A lliso n and B rook s (1 921 ). T hey con clu ded th at ch anges in the bon e w ere p ropo rtio na lto the ex ten t o f the in te rfe rence w ith norm al func tion , regard less o f h ow the d is tu rbance w aspro duced . F urthe r exp er im en ts to exam in e the t roph ic effec ts o f nerv e les io ns w erepe rfo rm ed by Tow er (193 7). In expe rim en ts up on lim bs pa raly sed by co rd iso latio n she fou nda trophy of b one w hen th e low er p art o f th e ne rvou s system w as in tac t b u t no t func tion ing .In show ing tha t bone w hich w as innerva ted bu t d isused w ou ld read ily change in d iam eter ,she co nc lu ded tha t the re w as no ev idence o f a d irec t troph ic fac to r in the nervo us sy stemin flu en c ing th e g row th of bon e.

W hereas it is accep ted by all inv estiga to rs tha t pa raly sis o f m uscle s leads to b one a troph y ,th ere is no agreem en t u pon the ch an ges in leng th w hich fo llow nerve d iv is ion . A rm strong(19 46) dene rva ted th e fo relim bs of e igh t imm atu re ra ts and fo und , fif te en w eeks late r , s ligh tshorten ing of the hum erus , and a little m ore sh orten in g of the rad ius. G ille sp ie (1 95 4) in th ek itten fo und no leng th changes in the lim b bones eig h t w eeks a fte r an ter io r ro o t s ec tion ,a ltho ugh th ere w as con side rab le bo ne atrop hy . These bo nes show ed a con side rab le red uc tionin w eig h t and rad iog raph ic d en sity com pared to the co n tro l lim b . A lthou gh the re w ere ve ryslig h t ch an ges in bon e q ua lity , the m a in e ffect o f the n e rve le sio n w as to p rod uce a d im inu tio nin the to ta l q uan tity o f b one rem ain ing . G ille sp ie s failu re to ob se rve p a ra ly tic leng then ingin the k itten w as prob ab ly due to the ear ly ske le ta l m atu rity o f th is an im al.

Changes in bon e fo rm occur in a ll m o to r de fec ts o f th e low er lim b . T hese a re m ostp ron oun ced in th e fem ur. Th e fem ora l neck re ta in s its in fan tile fo rm . T he positio n of va lgu san d an tev ersion is no t con fined to p o liom yelitis, b u t is comm on to a ll p a ra ly tic le sio ns, an dm ay a lso be seen in assoc ia tion w ith de lay in w a lk ing du e to m en ta l d efect. C hangesin th e ang le o f the fem ora l n eck w ill p rod uce variations in the leng th of th e fem ur on th escanog raph , bu t th ese va ria tio ns ap pea r to b e sligh t. T he low er fem oral ep ip hys is is a lso a ffec tedin m os t ch ild ren w ith a low er m oto r n eu ro ne le sion . T he de fec t is alm ost a lw ays on e of arela tive in crease in the g row th ra te on th e inn er sid e, w ith the con sequen t deve lopm en t o fa genuv alg um . Su ch a de fo rm ity m ig h t b e du e to a con trac tu re o f th e so ft tissues o n th e ou ter sideof th e kn ee, to the d efo rm in g effec t o f w e igh t bearing w ith ou t m u sc le sup port, o r to theva lgus po sition o f the h ip .

Changes in the sh ape of the tib ia a re sligh t and incons tan t. A secon da ry ro ta tiondefo rm ity occasio na lly appears , and is accen tu a ted by backw ard d isp lacem en t o f the la te ra lm alleo lus . T hese chan ges do no t in f luence m easu rem en ts o f tib ia l leng th . C h anges in thesh ape of the fo o t b ones a lso occur, e ithe r as a d irec t resu lt o f m usc le in activ ity , o r second aryto so ft-tissue con trac tu res p rodu cin g defo rm ity . T hese chang es m ay pro duce insign ifican te f fec ts upon fu nc tiona l lim b leng th .

CONCEPTS O F NERVE CONTROLThe c lin ica l and exp erim en ta l ev id en ce w h ich h as been pre sen ted an d d iscussed ind ica te s

tha t the no rm a l g row th an d m a in tenan ce of b one is dep en den t up on m usc le ac tiv ity . S e lectiv enerve les io ns h av e sh ow n tha t senso ry and sym pa the tic f ib re s do n o t in f luence the g row th o rstru ctu re o f bone . L esions of th e m o to r ne rves p rodu ce chang es w h ich m ust, in the end , bedue to the d is tu rbance o f lim b func tion w hich fo llow s m u sc le pa ra lys is . T h ere is no ev idenceto confirm the ex isten ce o f troph ic ne rve fib re s .

D e fec ts o f m u sc le ac tion m ig h t b e ex pected to a ffec t th e d ev e lopm en t o f th e sk e leto n inone of sev eral w ays . F rom the lo ss o f d irect m usc le ac tion , o r as a resu lt o f a d istu rbance ofwhole lim b fu nc tion , the re m ay be a m ech an ica l in f luence u pon the g row th o f the lim b .C hang es in tis sue tem pera tu re w ith in the lim b and in m usc le b loo d flow a lso o ccur . In the

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THE IN FLUENCE O F THE NERVOUS SY STEM UPON THE GROW TH OF BONES I 37late stages of the p araly tic lesion fib ro tic chang es in m u sc le m ay be assoc iated w ith th ere ta rda tion o f bon e grow th .

P e rio stea l n ew bone fo rm atio n is un doub ted ly d istu rb ed by a loss o f m usc le p u ll. A s are su lt, the pe rio steum becom es tig h tly b ound to the u nd erly ing bon e, an d th e osteo gen ictis sue is scan ty . M usc le m ark in gs a re less w ell deve loped . A ll these chang es can be ex p la inedby a d irec t lo s s o f m u sc le pu ll. In the ch ild m ost m uscle s a re attached to pe rio steum ra the rthan to bo ne itse lf, an d th e in flu ence of m u sc le loss is a d irec t and lo ca l o ne . W here the lo sso f m u sc le pu ll is con fin ed to a sm a ll a rea , lo ss o f bo ne bu lk is la rg ely confined to the sam ereg ion . T his is pa rticu la rly w e ll show n in the a tro phy o f th e tro chan te rs w h ich o ccurs in th epresence of a loca l m usc le para lys is. I t is le ss obv ious in the m usc les w h ich c lo the the sha ftso f long bones.

To th e fa ilu re o f the bon e to g row in d iam ete r m ust be add ed the lo ss o f m uch of itslam ella r struc tu re , atrop hy o f the co rtex , and en largem en t o f the m edu lla . Th ese ch an gesare p ro bab ly du e less to th e los s o f loca l m usc le p u ll than to the m ore gene ral d is tu rbanceof use w h ich is as soc ia ted w ith a pa ra ly tic lesio n . Th e pa ra ly sed lim b tends to b e sp a redfrom th e stra in s and stres se s w h ich are re spo nsib le fo r the m a in tenance o f fu ll b on estructure.

In a d iscus sion of the e ffects o f ne rv e les ions it m ust b e rem em bered tha t the g row th o fb one in leng th o ccurs m a in ly at the m e taphy sis , bu t sm a ll inc rem en ts occu r in th e bo nyep iphys is b y th e rep lacem en t o f a rticu la r ca rtilage . F rom a s tu dy of th e ex perim en tal an im alit is p robab le tha t the ep iphys is in para ly tic les ions d oes no t take part in the p rocess o fpara ly tic leng then ing , an d its g row th , ev en in the early s tages of a n erve les ion , is reduced .T he changes in to tal bo ne len g th the re fo re depend upon w he the r the in crease in d iaph ysia lg row th exceeds the depres sion of ep iphy sial activ ity .

I t w ou ld be reaso nab le to assum e tha t the new bon e fo rm ed by the m etaph ysis dep en dsupon m echan ica l fac to rs fo r its m ain tenance . U s ing th is a rgum ent, the d istu rbance o f g row thin leng th w hich is assoc ia ted w ith a p araly tic les ion depends , in th e en d , upo n th e sam e fac to rsas the d is tu rbance of g row th in bread th . T h ere is no reaso n w hy the m etaphys ial bon e sho u ldbe less sen sitive than the perios tea l bon e, o the r than in its iso la tion from the d irec t loss o fm usc le ac tion . A lthou gh it is cus tom ary to rega rd the g row th in leng th o fth e b one as depend en tp rim arily upon the actio n of the ep iphy sia l ca rtilage , the re can be no doub t th at th e speedof g row th in th is reg ion is a lso govern ed by the ra te o f new bon e fo rm ation in themetaphys is .

T he w eakness o f the m echan ical exp lan atio n of b one shorten ing lie s in its fa ilu re toexp lain the p henom enon of p ara ly tic leng th en ing . T he re is little d ifference be tw een thefun ctiona l s ta te o f th e severe ly para lysed lim b one m on th f rom the on se t o f th e d isease andth e fo llow ing y ea r . It is tem pting to rega rd the fib ro tic ch an ges in m usc le a s p lay in g a pa rt inth e dev elo pm en t o f lim b shorten ing . H ow eve r, th e lim b w ith con trac tu re s is n o m o re like lyto dev elo p seve re sh orten ing th an the lim b w ithou t (R ing 1 957 ). A lso it is c lear th at con trac tu re sof m usc le can deve lop in the early s tages of the d isease , w h en len g then ing is the ru le . Para ly ticleng th en ing canno t be d irec tly du e to th e m ech an ica l d is tu rb ance of func tion , and th is m ustlead to an exam ina tion of th e vascu la r facto rs .

T he to ta l b lood flow in the lim b a fte r p o liom ye litis is usu ally d istu rbed , bu t w hen th is isrela ted to the v o lum e o f the p ara lysed lim b the re is p robab ly n o sign ifican t d im in u tion oftissue b lo od flow at res t. H ow ever, as has been seen , the v asod ila tion response in the sk in issom etim es absen t, and the b lood flow in ac tiv ity m ust be reduced . O ne ofthe resu lts o f inactiv ityis a co ld leg , and th e tem pera tu re changes in sk in and m u sc le a f ter p o liom ye litis reco rded byT ro tt, N es line and G reen (1 958 ) have b een con firm ed in a sm a ll pe rso na l s er ie s o f pa tien tsus in g a s tee l/cons tan tan therm o coup le . It m ig h t seem reaso nab le to sugg est tha t th e sho rten ingof th e lim b m ay b e rela ted to th e low erin g o f tissu e tem peratu re a f ter m usc le pa ra lys is .However, hea tin g th e ep iphys ia l ca rtilage in th e experim en ta l an im al (R in g and L ee 1958 )

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13 8 P . A . R INGfails to p ro duce bone leng then in g . In fo u r ch ild ren w ith lim b shorten ing fo llow in g po liom ye litisthe ep iphy ses a roun d th e k nee w ere m a in ta ined a t a tem pera tu re o f 105 deg rees F ahrenhe itfo r th ree m on ths . N o chang e in th e ra te o f b one grow th occurred .

If the e ffec t o f p ara lysis is n o t d irec tly a the rm o-regu la to ry o ne , it is a t lea st p ossib leth at som e v ascu la r shu n t m ay p ro duce chang es in lim b leng th . In an expe rim en ta l s tud y o fbo ne rare fac tio n fo llow ing ten o tom y of th e tendo ca lcan eus o r p la ste r imm obilis atio n G eiserand T rue ta (19 58) show ed th at bon e ra re fac tion is in tim a tely assoc iated w ith h ype raem ia.This o bserv ation p resen ts a log ica l exp lan ation of the para ly tic leng then ing o f bon es w hichoccu rs soon a fter a m o to r ne rve le sion , o r in deed , du rin g any p eriod ofin activ ity . L eng th en ingof b ones in the ea rly stages o f po liom yelitis is like the leng then ing w hich occurs in the earlystages o f th e tube rcu lous h ip -b o th a re due to b one h ype raem ia .

T he hype raem ia o f d isu se is tem pora ry , and o ve rg row th o f bo ne ceases w h en its b loodsu pp ly retu rn s to no rm al o r sub norm a l lim its . M igh t th e sub seq uen t sho rten in g of b one bed irec tly re lated to bon e isch aem ia? It h as ye t to be show n tha t the m e taphy sis in the pa ra lysedlim b suffe rs a s ign if ican t d im inu tio n in its b lood supp ly , and if it d oes, th at th is ischaem ia isth e cause o f th e depres sion of m e tap hysia l activ ity ra the r than o ne of th e effec ts o f a d im inu tio nin the ra te o f bo ne g row th . Th e m ech an ica l in fluen ces w hich regu la te the decom position o fb one canno t be ligh tly d isreg arded , and it ap pea rs poss ib le th at b o th v ascu la r and m echan ica lch an ges p lay a pa rt in th e fin al d is tu rbance of bon e grow th .

Tw o fac to rs thu s seem to b e of sig n if icance in the chan ges o f bo ne g row th w hich fo llowp ara ly tic le sio ns : I ) The hype raem ia o f d isuse w hich pro duces tem porary s tim u la tion o f th era te o f bon e g row th and re su lts in the p henom enon o f pa raly tic leng then in g ; 2 ) the ch an gesin lim b fun ctio n w h ich re su lt f rom para ly sis an d d isu se. Th ese m echan ica l changes a reas soc ia ted w ith a re tarda tion of g row th in d iam e te r and leng th .

SUMMARY1 . A n expe rim en ta l s tu dy o f th e e ffects o f ne rve and m uscle le sio ns up on the g row th of bon eh as been m ade . In each case an im a ls w ere su b jected to un ilate ral les ions in the h ind lim b ,th e o the r lim b se rv ing as a con tro l. T he grow th of the tib ia w as m easu red b y ca lcu latin gth e d iffe rence b e tw een th e leng th o f the b on e on a rad iog raph at the b eg inn in g of theexp erim en t and the leng th o f the d ried bo ne a fter nec rop sy . Th e w eig h ts o f the d ried b oneswe re compared.2 . In the young rab b it s im p le exp osu re o f the comm on pe ro neal ne rv e, o r d iv is ion o f thesu ra l n erve , p ro duced no ch ange in the g row th rate o f the tib ia . D iv ision o f b o th pe ro nea lnerves , p roduc ing para ly sis o f the m usc les be low the knee , led to leng th en in g of the affec tedtib ia , and th is leng th en ing pe rsis ted u n til m a tu rity seve ra l m on ths late r . A s im ila r leng then ingw as seen afte r d iv is ion o f the ten don s a ro und the ank le. In sp ite o f th is len g then ing the tib iao n the sid e o f the o pe ra tion w as alm ost a lw ay s lig h te r than its fellow .3 . In the p uppy d iv ision o f the an te r io r n erve ro o ts sup p ly in g the h ind lim b pro du ced asig n ifican t leng th en ing of the tib ia o f th e a ffec ted lim b th ree m on ths a fte r o pe ra tion . N osig n ifican t chang es in lim b leng th occu rred a fte r lum bar sym pa thec tom y in th e puppy .4 . T he s ig n if icance o f these exp erim en ta l ne rve le sions has b een con side red toge the r w ithrecen t obse rva tions u pon the g row th of b one in the p re sence o f low er m oto r n eu ro ne le sion sin the ch ild . F rom th is ana lysis it is sugg es ted tha t the in itia l e ffect o f p ara lysis is to p rodu celeng th en ing o f the a ffected b one . T h is len g then in g is p ro bab ly du e to the hy pe raem ia o fd isuse . In the p re sen ce o f pe rsis ten t p ara lysis the g row th o f the lim b is u ltim a te ly depre ssed .Th is d ep re ssio n is ra re ly seen in the exp erim en ta l an im a l because th e g row in g pe rio d isre la tive ly sh ort. T h e pos sib le causes o f th is s econ da ry dep re ssion of b one g row th havebe en c ons ide re d.

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THE IN FLUENCE OF THE NERVOUS SY STEM UPON THE GROWTH OF BONES 13 9

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GULL ICK SON , G ., Jun ., KUBICEK , W . G ., and KOTTKE , F . J . (1951): E ffec ts ofS tim ulation o f Lum bar S ym pa th eticNerves on Long itu d ina l G row th in D og s . Federa t ion Proceedings , 10 , 56 .

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HARR IS , R . I., and MCDONA LD , J . L . (1 936 ): T he E ffec t o f L um bar S ym pa thectom y upon the G row th of L egsPara lyzed by A nte rio r P oliom yelitis . Journal o f B on e and Join t S urg erj, 18, 3 5 .

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th e A cu te P ha se o f P oliom ye lit is . Acta M edic a S can din av ica , 114 , 565.MCPHER SON , A ., and KESSEL , A . W . L . (1955 ): A symm etry ofB lo od -flow in th e F ee t in A nte rio r P oliom ye lit is .

C lin ica l Science, 14 , 361.MCPHER SON , A ., and KES SEL , A . W . L . (1956) : D uration o f V aso d ila ta tio n a fte r L um ba r Sym pa th ec tom y.

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Sym pa the tic T ru nk in th e D og . A natom ical Record , 113 , 421.MEYER , H . (1 867 ): D ie A rch ite c tu r de r S pong io sa . A rch iv f# {2 52 }r na tom ic, Ph sio log ie und W issensc /za ftlic /ze

Medicin , 615 .MURRAY , P . D . F ., and HUXLEY , J. S . (1925): S elf-d if ferentiation in the G rafted L im b -bud ofth e C hick . Journal

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P ro ceedin gs o f the R oyal S oc iety o f M ed icine (Sec t ion of O rtho paed ics ), 26 , 429 .VOL . 43 B , NO . 1, FEBRUARY 1961


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14 0 P . A . R INGPOTTOR F , J . L . (1 916 ): An Exper im ental S tud y of B on e G row th in the Dog. A na tom ica l Record , 10, 2 34.RATLIFF , A . H . C . (1959 ): T he Short L eg in P o liom yelitis . Jou rna l o fB one an dJo in t Su rge ry, 41-B , 56 .RING , P . A . (1957 ): Sh orten ing and P ara lysis in P o liom yelitis. Lancet , ii, 9 80 .RING , P . A . (1 95 8): P ro gn os is of L im b Inequa lity F o llow ing P ara ly tic P o liom yelitis. Lancet , ii, 1 ,3 06 .R ING , P . A ., and LEE , J . (195 8): T he E ffec t o f H ea t u pon the G row th of B on e. Jou rna l o f Pa tho log y and

B ac ter io lo gy, 75 , 405 .RING . P . A ., and WARD , B . C . H . (1958 ): Para ly tic B one L eng then ing Fo llow in g Po liom yelitis . Lancet , ii, 551.Ross , J . P . (19 46): T he Surge ry of A rte ria l D isease and In ju ry . B rit is h M ed ic al Jo ur na l, i, I.RossI , F. ( 1932 ) : L in nervaz io ne de l m ido llo osseo . A rch ivio I ta l ian o di A natom ia e di Em brio lo g ia , 29 , 539 .SA NCH IS PERP INA , V . (1955 ): L # {233 }nerv ation sym path iq ue d u m em bre in f#{2 33}rieu r. L yon C h iru rgical , 50 , 526 .SHA RRARD , W . J. W . (1 9 59 ) : P erso na l c om mu nica tion .SM I TH , E. , RO S ENBLATT , P ., and L IM AURO , A . B . (1949): T he R ole of the Sym pa th etic N ervous S ystem in Acute

Pol iomyel i t i s . J ou rn al o fP ed ia tric s, 34 , 1 .ST INCHF IELD , A . J., RE IDY , J . A ., and BARR , J . S . (1949): P red ict ion o fUn equalG row th o fth e L ow er E xtrem ities

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British M ed ica l Jou rna l, ii , 770 .TOWER , S . S . (1937 ): T ro ph ic C on tro l o f N on -n e rvou s T is su es b y the N ervo us Sys tem . A S tu dy of M usc le

an d Bon e Inn erv a ted from an Iso lated an d Q uiescen t R eg ion of S p ina l C ord . Journal o f C om pa rativeN eu rology , 6 7 , 241 .

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Z I NN , C . J., and GR IFFITH , J . Q ., Jun . (194 1): E ffec t o fS ym pa thec tom y upon B lo od Supp ly of B one . Proceedingso f the Socie ty fo r E xp erim en ta l B io lo gy and M edic ine , 46 , 311 .

THE JO URNA L OF BON E AND JO IN T SU RG ERY

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