Acta BioI. llltng.

, 14 (2), 145-155 (1963)

FUNCTION OF HETEROTOPIC SPINAL CORD
SEGMENTS INVESTIGATED IN THE CHICK
K. STIlAZN leI(Y
IlEI'.\IITMENT 01' ANATOMY. IIISTO!.O(;Y ANIl EMIlIIYU!.O(;Y. MEIlICA!. lINIVEIISITY. I'I::CS
(IIEAD: .1. SZENTA(;O'l'IIAI)
(Heeeived May 15, 1963)
Synopsis
In rhick cmbryos (Hamburger and Hamilton stages 15 and 16) the previously
extirpated braehiul cord segments were replaced by brachial, thoracic, or lumbo-
!illerul coni t nken from otheremhryos ofthesame age. Innervation nnd func-
lion of thl: willJ.:s illllcrvllled by the grafted cord segments were investigated after
hllldlillJ.:. 'I'h,' brarhial :legmcnls of another chick can substitute for the extirpated
hrarhilll sCJ.:Illr.nl in every respect. Normul innervntion and function ofthe wings had
developed in such eases. Although the nerves of grafled thoracic segments innervated
th(' which to pain, Ihe nlllsculuture hecullle completely atrophic
alld I he join t s an ky lot ie. II ypertroph y developed in the dorsal grey columns of the
J.:rufted slJ.:menls, whereas Ilw molor column exhihited certain siglls of distrophy.
Wings inllervated by lwterotopie lumbo-saeral segmcnts showed moderate motility
in lhl' :lhollidcr. synehronizl,d with the movcments of the legs 011 the same
:lidl. III spite of wI,11 Jeveloprd IInJ nornwlly inncrvllted musculature 110 coordinated
movclnenls in Iht' olht'rjoints (:ould be ohserved. The possihle mcchunisms underlying
the immohility or Ihe distal joints lind the synchronizcd movements in the shonlder
nr('
Introduction
A lIumlll'r ordata suppurt the that coordinated limb move-
ments can he eontrolled only by the apprupriate limb innervating segments
of the spinal eonl in till' newl [1,3,10,15]. Supernumerary limbs, both in
amphihiaand hirds,illlwrvatcdbytrunksegments remained motionless [9,
unless a sillgl<' hraneh from the brachial plexus, howeversmall,contributed to
lhe limh 14.]. recently direct experimental evidence was
hySZEKELY 113] for the ofa specific segmental apparatus
in the cord atlimb I('vel, capableofelicitingcoordinated function oftheinner-
vated limh. By transplanting brachial or lumbo-sacral segmental groups into
the plae(' ofthe previously excised thoracicsegmentsandadditionallygrafting
Illipernumerary limbs at the same level, he succeeded in demonstrating that
hcsides perfectly coordinated stepping movements the participation of the
Ilupernumerary limbs in the stepping pattern was characteristic ofthe natun'
of the limh innervatingheterotopiccord segments. As alloftheseexperiments
were performed on amphibia,itseemed promisingtoextend theinvestigations
tu birds whose moreelaboratelocomotion patternsand centralnervoussystem
Aria RWl. HuRf,. 14. /963
14<1 K. STIIAZNICK'
of histologically higher diffl'rcnl.iation might contrihlltt' to tht, I'rol>lt'm of th('
funetioll of ht'l('rot0l'ic i-ipinal cortI i-i('gm('llts. WI' altt'mpl.l'd, tht'rcfoTt" to
replace th(' wing innt'rvatillg brachial s('gmt'nls hy thoracic and lumho-sacral
s('gm(,lll,,1 groups in chicken t'mbryos and to invt'stigatt' th(' motility of such
wings afLt'r hatching. An account of t'xl'('I"imt'lIts is I'Tt'st'nLcd ill this
pap('T. 1\11 abslrat't or lhis l'''l'l'r has ht't'll I'uhli:,dll'd ('ls('wht'IT 112).
Method
Till' W('I'l' pl'rfllrlllt'd llll lhl' elllbryo... of illbl'('d ... Irail)s llf l\\ll loeal vlIrictil''''
or fowl (IIIIII/-(ariall Yellllw alld Spl'ekled). '1'111' ('llIbryo" \\'I'!'I' 0pl'rall'd 111'011 Oil Ihl' third du)
or inl'lIhalioll ill IS ulld 1(, llr IIAMIII'IC(;EIC alld IIAMIL'I'ON 1111, Thl' Ihllraeie cord
or Ill(' dOllor wI'n' I.y tltc' aid of tltc' (''''('ll'OIlIUp:III'lil' vihralill/-( lI!'l'dlc lSI,
Ihl'lI cal'('fllily clculH'd rrolll dlOl'(l;1 alld IlIylllollll' lIIall'rial alld arll'r I ill Ihl' lIccep
lllr I'g/-(, riltcd illio I It I' plncc llf I Ill' 1'!'I'violl"l)' n'llIovI'd I'radlial (I () III I C)) cord
III another grollp of thl' (2:1 10 2lJ) ... IU'!'I' grafted illto
the SUIIW plac'l', To ('olllro! Ihe illfillellf'c' llf IIII' opl'ralioll it ... c,]f 011 willp: flllldioll, the
hrurhilll (.1' 111'0 crtlhr)'os II'('!'I' illlerchnllg('d ill Ih(' Ihird ;..(l'Hllp of n:I'('ri,llcnl:J1
1Il1illlals. Frol!l a Illilli llf 22() opl'l':lI('d t'II1hrylls wc' ill /.!!'Ilillg 17 via hi!' hntchl'd
rhic'ks, rrolll which 1/. hoI'(' t IlOr"c'i(', II 1IIIIIJlflsa('J',,1 S('gllll'1l1;; Oil I hI' pla('(' of I ht, 1't'llIovec!
hrachial wltil(' 2 allilllais helollp:t,cI 10 Ihl' c'olllrol group. Aftl'r halchillp:, Ihc' 1Il1illlal ...
kcpt alivl' frnlll 2 10 :; 1I1fl1l1 h.... Dllrill/-( I pc'riocl 11101 iii I yoI' I ht' was ilhscr\"
I,d ('olltillilously 1I11c1 till' IIt'rt' rt't'orclt'd I,y Cilll'llIall'llgruphy. Aftl'r
sarrifirlllion lht, anilllal" wt'rl' fixld ill 1'0 r Illfll 1:1,
(ullt'rvalion alld of lhl' II ('n' slilcliell parlly hy allatomical
dissl'c'lioll ullci partly in lIIi('J'osl'opic:d wil h Bic'lsl'lwwsky's lIll'thod.
l
Thc' hisl ologica' ip:a I ion of I hI' /-(ra rt I'd ('ord S(';..(IIH'1I1 hu,.. h('c'lI Iwrforllll'c1 ill Bi('I
srhowsky sluilll'cI
I
Tht, ge'llt'nd jH'ha\'iour alld growL II of I he' 0l't'ra I"d "hicks disn'gardin:-:
th(' willg!'! \Va!'! Ilormal ill Ihe' llIajorily of I'lU'WS, HI't'ollslruclioll of tht,
v('TLt'hral eO\UII1I1 alld fllsion of Ih., grafLt'd :-;l'gJlll'Jlls wilh lht' J'('llIaJlIIJlg cut
Pi/!.. 1. findillg ('olll'('rllillg all allilllal ill whil'h thl' hral'hial s('gillellls hud uel'lI
hl'lwl't'lI I WO "III :'>I orlllal hral'lJilll pl(,:'( hud dl'v('lopt'r1 frolll t ht' 12th
anrl I:-llh ('I'rv;('nl alld frn II I Iht' Ihora(';,' s"l!:lIlrlll. A slrollg hralld. 1'1'0111 II... 211d
t horlleic' s('l!:lIlt'nl ron I rihu I 10 I hr pltx us for ilia I iOIl. This l'hi"k kepI uliv(' for S 1Il0nths,
Filf,. 2. filldill/-( ill lllflrul'il' SI'gllll'lIts graftt'd inlo thl' plnl'l' or hruchiul sc'/.:mcnls.
The spillul ('onl d('vl'lop!'d lIorulully. Th(, hruchial plt'xlIs llll t hi' from thl'
lith, 12th, Ulh el'rvil'ul ulltl frolll lIlt' lSi Ihor,ll'il' S('I!:II\1'III. while, 011 tilt' 11'1'1 from tht'
121h, 131h ccrvit'nl ulld frolll lhl' nlld 211c1 lhoruci(' Thl' chick sllcrificcd 6
1I1'1t'r hatthill!t
{-'ilf,. 3. fillding ill Jlllllho-sal'ral segll)('lIls gl'aftl'd illio 'hr ,,111('(' of "rachilll St'l!:llll'nts.
Thl' sJlillll1 l'Onlllt'velopl't1l1llrlllally ulld tht, glial (arrow) was Jln'served,
Thl' "I'Hlldw!' of Ihe hral'hial t'llllrp:t'tJ. as ill lIorllllll fro 1I I fOllr SI'I!IIIt'llts. Th('
t'hiek al I ht' lI;..(r of S 1110111 lis
4 1II1l1 .S. 1)('IIIOIISII'II11' Ihe c1iff(,rt'lI('(' IWIWt'l'!1 Iht' 1101'111111 (Fig. ,n III1UII !tl'lIfll'cl 5)
thorac'ie eon! of Ihe Slllllt' IIl1imal. Thl' dOl'sal horll alld lhl' illl!'r1111'dillll' of thl' I!:rufted
sel-(llIelllS I'ullll'g!'d IIlld a rl'lluf'I;oll ill lIumh!'1' 01' vl'lIlral 1IJ1J1l'nr!<
to hllvl' tnk('lI plm'r. MII;..(lIifirulioll 6S -:
Fi/{s. (, lIud 7. Normlll Illotor 1'1l11-platt's (arrow) havI' c1cvcloJll'tJ ill the willg musclI!nture,
iIlIH'rvlltl'<! hy p;raftl'd IlI1Ilbo-sanal
./rlll 1Ii,,'. 1/1I111:' 1I. I%,'j
146 K. STnAZN1CKY
ends of the spinal cord was also complete, so that no sign indicated the sitt
of operation. The function of the legs in 5 animals was more or less damagec].
these chicks were mostly sitting on their belly and unable to perform normal
locomotion. Small protrusions and deviations were found on their vertrebral
column ana the grafted segments were connected with the lower spinal coru
through a somewhat narrowed region. No immunological reaction was dctecteel
even in animals which were kept alive for the longest period. The fo})owing
characteristics were found in the several experimental groups.
1. In the two successfully hatched cases, ill which brachial
had heen exchanged between two embryos, the function of the wmgs
Fig. 8. The of II chick wilh l!wrncic sep;IIII'nls grnfted inlo lhe pln('l' of Ihe hruchinl ones.
The musculature of the free extremily hilS completely disappt'nr'd, Il:l VI' for a thin hundle at
the place of the triceps. The median nerve (fulliinc) nnd th' Tudillllll'rv' (pnrlly hrokl'n line)
follow the norlllnl COUTS' of the wing IlI'TVI'S
indistinguishahle from that in normal animals (Fig. 10), The disst'etion findings
revealed t:omplete fusion of the ends and normal plexus formation (Fig. 1).
The arhorization pattern of the pt'Til'ht'Tal nerves and the' mllsndaturl' of the
wings, in as much ;IS it coulel he traced undrr c\i!'sccting microscope, was
exactly similar to those of an intact animal. Formation (If rpgular e'IHI-platcs
was seen in the impregnated wing muscle's. 1Il'tt'rotopic hrachial sqrmf'nts
can, thcrpfore, for the normal onps in eVl'ry Tespl"CL
2. When tht brachial segments hall L''n rl'Jl1a(,pd lly thl' Ihoracic
segments, the wings [emain't! pl'rf'pctly lJloLionll'ss. Thl' joints l)('carn' anky-
lotic and the wings were slightly relardl'd in growlh. '1'ht, chich wt'rt' IInahlc
to stand up when they haprll'nt'd to fallon tht sid' Of to 1)(' t1lml'd upside
down, although locomotion by thr IPgs was normal oLhpf\visr. Thl' animals
exhibited a remarkabll' sl'nsitivity to painful stimulation of Ihe wings, hut
.'Irln lIiol. 1/""/1, /<I. 196:1
I

j
147 IIE'I'EIlO'l'OP/C .'iI'INAI. COIlIl SE(;MENTS IN TilE CIIICK
they could not localizl' th(' ('xact placI' of stimulation. The general locomotion
of th(' animals was nurmal, save for some slight uncertainty in
maintaining the 1'(lllilihriIlJn during cl'rtain strained as for
Jumplllg.
!\(:(:ordillg to findings, the fusiun hdwcen the and
ortholopic sq!;nll'nts was compl(,te (Fig. 2). Macroscopic enlargement
failed to d"vdop an(l the nerves formed a p[l'XUS before entering the wings.
In spit(, of till' faet that five primordial thoracic segments had to be trans-
planted in order to fil.l in the gap left by brachial segments,
four s('gm('nts devdul'ed from the graftecl cord, and the innervation of the
wing, like ill intaet animals, from the 13 to Hlth spinal nerves including
a small branch from the 12th nerve. The nerves, however, w('re thin,
hardly the thickness of normal thoracic nerves. The spinal ganglia,
<IS compared with the normal thoracic ganglia, were larger in size, owing to
their rdativ('ly overloaded I'cril'heric fitld, but they did not reach the !'ii:l.e of
1I0rmai hrachial ganglia. Tht' lH'ripheric arborization pattern was suggestive
Hf the normal wing (Fig. 8). Th(' radial and median nerve could be well recog-
nized; flU sidl' hranchcl;, howI'ver, WCft' fOlln(1 in macroscopical preparation,
owing, appanntly, to lack of musculature. This corroborates PlATT'S [9]
('urlier observation that the arhurization is determined by the struc-
ture of the inllervated The histological pictures of the heterotopic seg-
ments revealed an enlargement, especially in the head of the posterior column,
(,(Illtaining more neurons than orthotopic thoracic segments (Figs. 4, and 5).
Considerable hypertrophy waS found also in the intermediate zone. The numher
of in the anterior eolumn, on the other hand, was not increascc.I.
This finding is in accordancc with HAMBURGER'S observations [7], who has
(It'monstratetl that motor eulumn of an thoracic spinal cord
tloes not change its size in eontrast to the sen!'ioric onc which shows a clear
in size to the peripheric load.
The mWicldature of such wings was Htrophied. While the muscles of the
although much smaller in !'iize, more or less persisted no muscle was
found on free extremity, except for a small distrophic muscle bundle in
place of trieeps (Fig. 8). This bundle was present in every case.
3. most interesting result of the experiments was the establishment
of the function of wings innervated by heterotopic lumbo-sacral segments.
A peculiar motility had. developed which occurred always parallel with move-
,"ents of the When walking, the animals raised and adduced the wing to
rhe shoulder joint (Fig. 12) exactly in time with the step of the leg on the same
I<itle. This movement of wing was of little excursion, about in the 15 to 20
t1t'gree range. Although the musculature of the free extremity was fairly
well maintained and normally innervated (Figs. 6 and 7), the joints were
mobile, no movements occurred in the other joints or they were so small as
Aria IJWl. flunK' U. 1963
14H K, STHAZNICK \
to escape dl'tection. Thl' wings n'mailll'd ('oJnpktely rnotionll'ss lIndl'r all
circumstances that Idicit thl' charactl'ristie wing n'f'Il'xI'S in intacL animals, l
Letting rail down a chick provokts rapid r1aLl\'l"ing of IIII' wings, or t
aroulld the longitlldinal axis an t'xtl'nsion of Ihl' wing 011 tht, sid( tllrned \
downwards and a rJI'xion and addllclion on tllf' ollt.'r, !\Jon!' of th('st' n'sllllllsl'S
appearl'd ill th(' opl'rall'd anima Is (Fig, J I). I t was, Oil I It I' oLIll'r Itand, ill I('f('sl
rr
\
ViJ!,. 9. '1'11(' willp: or 11 (lti('k witlt lll,"II1l-sanal "('P:"I1Il..; p:rul't('d i"to th( plan' 01' ItradJiul
I
01' Iltl' Itav!' d'V(,lol'(d. fwdpl,,'ral "ollrsl' alld :Irltoril'.alioll I'utlem
\
of th.' lII!'diulI (flllllilll') lIlld lit.' rndillllll'rvl's (1lrllkl'lI lilll') lilT silllilar 10 I ill IIII' Ilol"lllllll)'
i,,"t'I'\'lIl1'd
ing to 1I0tl' that whl'1I Ihl' ehil'k \las Ill'('king alld IWllding c!o\\n for grains
on Ihe ground, all ITlO"I'IllI'IIIS involving 111(' hip joinl \\\'1"(' a('('(lIIlIHlIJi"d by
thp motions of thp willgs dl'snilll'd ,,110\'1'. Frolll lit .. silldy of 1111' fllnC'lion of
the wings thl' g('I\Nal impn'ssion was gaillld thaI lilt' innl'f\'ating 1111111.0-
sacral !owgmellts on tIl\' plal'" of till' hra('ltial sl'gnll'nls did nol real'l 10 ('pnlral
iml'uhws "ddrt'ssl'd 10 tltl' hru('hiul ('ord, hUI \\('n' sl'l into \\ork Silllllllul\I't1l1s1y
with thl' orthotopie lumho-s,uTal l'wg III ('11 Is.
Disspetloll sludi('s f('v('all'c! IYl'i('al IlIlIlho-san,,1 curd in Ih,' hra('hiul
n'gion with lll(' 1']lHnH'l"ristil' glial i'i\\'('lIillg ill Ih(' rhomhoidal Silllli'i (Fig. 3).
[<,iJ!,s. to. IT alld 12. Show 1110\('1111'111111101 I'll> t 111'1' ill ('hid,s \\ilh "nll'hial sl'illnl
l'('pllll'('r1 jl'ally ill I'arlv t'(lIhryolli(' Fig, to. Th,' hl'a('hi:d hn\'(' 11l'('1I
l'l'pIHf'(d hy lilt' lIIII(' or nllolhl'r ('llIhryo or silllilal' CharnC'l.'l'islil' rt'fh'xI's
"POII qllic'k dowllward 11111\'('1111'111 of till' halld, Fig, II. I.lIl1d,o-sal'l':r1 t'lIl'd illio Ihl'
I'lac'(" or 1111' hrll("hial AltllOllp:h 1IIIIS"I"s :11'<' \\1'11 illlll'I'\'III"d alld I'nirly \\("11
tl.'vdoptll. ,hl'l'(' is 110 1'I'1'I.'x dO\\II\1'lIroi IllO\"'II11'1I1 of Ih(' halld. 12, Silllilar
('al'(' as ill Fig, /I. wilh l'haral'tl'risli .. willg so III ('\,}I:II I'xll'lIdl'd IIl1d IIhdll,'('d, Sillllll-
with ('adl sl('p lht, ipsilllll'l':r1 is ('1"lIdy t, ... l(llIlt-d Hlld "lllllIl'('d
(('I J lIi,,1 11"".'_. II. (tJ",]
150 K. STnAZNICKY
From the seven primordial lumbosacral segments transplanted four corel
segments developed and four nerVI:S emerged which formed a plexus befon'
entering the wing. The rearrangement of the hetcrotopic spinal cord in thl'
number of segments both of lumbo-sacral and thoracic origin, was apparently
influenced by the number of the neighbouring myotomes, ns alreHcly showll
by DETWILER [2] in the newt. The four plexus forming spinal nerves wen'
mostly thicker than those of the heterotopic thoracic segments, hut did nol
reach the thickness of the normal hrachial ncrveR. Although the lumbar
enlargement was present in brachial position as well, the histological examina-
tions showed considerable loss of motoncurons as eompa,red with the f10rmal
lumbo-sacral segments. The general histological appearenee of the
left did not differ from that of the normal ones ill any part of the spinal cord.
The branching pattern of th(: peripheral nerves, as far as it eould be tract'11
under dissection microflcope, was perfectly regular (Fig.9) although tile'
nerves were considerably smaller than in the normally innerva Led wing:o;.
The same held true for the muscular apparatus of theRe wingR: all musclt,:o;
were wen developed, but their siz;e did not reach, in general, that of normal
wing muscles.
Discussion
I
As this investigation aims mainly at Rtullying the function or hCLerotopil' I
cord segments of different origin, the morphological findings arl' only briefJ)'
outlined, a more detailed discussion of myotrophic aspects heing rt'sl'rved for
a following paper.
From the results the unequivocal conclusion can 1)(' drawn, that clear
differences prevail in the functional capacity of diffl:r('nt spinal cord segments,
which arc determined already in early emhryonic life (HamhllTger-Hamilton
stages 15 and 16). Foreign brachial cord can fluhRtitute for the hORt's own
brachial cord in every respect. As to the function of connccting higher and
lower levels of the eNS, all segments Recm to he t. Thoracic segmcnts
arc unable to perform whatever movement in the wing, which supports the
several data available in literature [1., 10, 11.]. The axons of thoracic moto-
neurons for some unknown reason cannot cRtablish d'fectivl' myoneural junc-
tion with wing musculature, which then'forl' soon ulldl'rgoeR compldt' atrophy,
just as in the experimentally induced nerveless limbs of the chick [6]. Sensory
nerves invading the wing can, on the other hand, promote pain and Loueh
sensitivity. This difference is clearly represent('d hy thl' distrophic motor
column and the hypertrophic: sensory columns in the histologieal pielUfl's of
the heterotopic thoracic scgmcn ls.
Wings innervated hy heterotopic lumbo-sacral sl'gmellts ar(' ahle to
produce, if not complete, still eOllsiderahle movement, synchronollRly with the
Acla I1ial. J/""g. H. 1963
151 II ETEI\()TOI'IC SPINA l. conI) SE(;M ENTS IN TilE CHICK
movpnu'nls of ipsilaLerallegs. There is no satisfactory explanation, as yet,
to aCCOUll Lfor the movcmen t in the otherwi-ie normally developed
and lH'rfpctly wings. SZEKELY l13] has rcported a similar obscr-
vatioll ill thl' nl'wt, according to which in forl'limhs innervated by lumbo-
sacral spgml'llts, although showing wdl coordinated stepping movements,
the l'Jhow moves only to that limited extent which can be in the knee
of normal hindlimbs. But the aclaptation of inappropriate segmental apparatus
to the task of moving a limh is much better in newt. It might well be that
some highn plastil,ity or the segmental apparatus at the moment of the trans-
plantation accounts for this difference. It must be considered, however, that
the gross anatomical d iffncnce and consequently also the requirements of
building up a highly specific nervous apparatus for coordinated limb move-
ment arc smal\n a newt fore- and hindlimb, than between wing and
of the (:hick. This consideration iil supported by the fact that both in the
newt and the chick the in a limb, innervated by the inappropriate
segmental apparatus, are hetter preserved in the proximal joints and progres-
sively 1m,s in distal ones, where the divergence betwecn the two limbs becomes
naturally larga and particularly explicit wing and leg.
Another oh1'lervatiun some is the perfectly synchro-
nouS movement of the wings with the legs on the same side (Fig. 12). In similar
"xperiml'nts carried out on newts, SZEKELY [13] could ooserv,: but an inclina-
tion to perform by the forelimbs movements synchronized with those of the
hindlimbs, when the lumbo-sacral segments were substituted for the brachial
He suggested that the hindlimb character of the lumbo-sacral cord
might hav!' been not completely determined at the time of transplantation,
lind the grafted segment1'l might have more or less adapted themselves to their
nt'W situation, resulting in forelimb trends of their function. Literature dealing
with embryonic determination phenomena includes a large body of evidence
that emhryunie differentiation of most tissues, in general, is earlier
,ll>tcrmilled in the chick than in the newt. The dissimilarity between these and
Szekely's findings can, thl:rcforc, be explained by assuming that the lumbo-
"aaal cord, heing in the chick definitely determined at the time of transplan-
tlltion, can maintain the hindlimb character in its function. Perfect synchro-
of movcments in both limbs and the fact that the wings remain motion-
under circumstanccs that in normal animals evoke wing responses, suggest
furthermore that the grafted lumbo-sacral segments can somehow select from
impulses descending from higher centres to answer only to those which are
nd.lressed to the normal lumbo-sacral segments. Trying to interpret such
.. responses of the grafted segments, one may think of two alternatives:
Ii) The motoneurons of the grafted and the normal lumbo-sacral segments
,"ight establish connections with exactly the same descending neuron systems,
11I1l1 these specific neuronal connections might rise to the synchronous move-
Ada BU>l. Hu"t;. 14. 1963
152 K. STHAZNrCK\
ments 1I1 thr ('xtremities. Thl' ('stahlishrn('nt of lh(' sp('cific ('onll('('lions mighl
beregulated bysomespccificI'rojl('rlil's,elairn('d by WEISS alld SI'EHHY 115, 11]
to exist in thp neurons, which would allow lh(' dl'v('lopm('lIl of
synaps{'s only betweensets ofcl'rtaill particular IIl'urOIlS havill/1: gailll'd during
nl'urogelll'sis qualitative sjwcificalioll to match racll oliln. (ii) Tltl' allnllativl' >
is that('('rlain parls oflhl' eNS have tlt(, capacity 10 allaly:-,;(' difTnl'll1 impulse
and answer only 10 appropriatc' OIl('S, as slI/1:g('sll,d ill ('arlier l
works from this IltJ.]. Th(, limb sl'gnll'lIls ofIII(' spillal c'ord might
have somt' properly of this kind, probably Oil thl' groullds ora pl,t'uliar slrllc-
lund arrangemcntofneuronsill th(' spgm('lllal apilaraills. Furlhl'!'('''I'C'rilll('nts
art' n('rrssary to analyze lh('st, p!'oblt-ms.
H1':Io'EH I
1. Ih:TWIl.I':H, H. S. (1920) Expt'riml'lIt Oil thl' trallsplalltatioll or lilllb!'> ill .III/MYHIII/Ill. Thl' l
rOrllllllioll or lIerv(' alld Ihl' rlllll'lioll or Ihl' limh. ,I. /';.1/1. ZrllJ/. :1I, 117 170.
2. B. S. Expl'rilllf'lIlal !'>llldy Ill' !'>pillul IIl'rvl' sq{rlll'lIlalioll ill
wilh l'I,rrrt'II('c' 10 till' plllrisl'p;IIIt'lIlal ('olllrihlltioll III Iht' hra('hi.d pln,lIs../. 1'.'-''11. /'llul .
67, .195
:1. Ih:TWILEIl, H. S. (19:l6) Ncurot'III"rylllllgy ...\11 (''(III'rillwlIl,d study. \C'I' '\ ode Mill'-
.11i 1111II.
i/.. H. S. CAHI'ENTEIt, H. I.. (llJ2
1
All "'III'rilllc'lIlal siudy or IIH' 1I1l'1'laalli,,1II (,r
c'oordillnlrd 1II0vrlll('uis ill ht'l('rolopil' IiIII"s. .1. (;/J/I'/', ."V'III .. 47, 11.27 ,1,1.7.
S. I)ltl
'
RHY II. V. I\. vil>ralillg 1Il'('dll' a IIlil'l'llsurgi('ul illslrllulIIII. S('i(,,,(,. 93,
263 2M.
6. EASTLICK, II. I.. Slllllil's Oil Irall"l'lalllt'd l'III1>I')'IIUil' lirllh" or Ih. ('ltil!-. Till' dl\IllIp-
II1l'II1 or'"llsell' ill 1II'I'V'('SS ulld ill illlll'rYall'd gran!'>.. /. 1:.lp. Z(;ol.. 27 19.
7. IIAMllliIlGEH, V. (19.19) MotuI' alld sPllsory hYl'pl'l'lasiu rollo\ljllg li'""'''I111 Il'allsl'lulalioll
ill ('hil'k embryos. Zool.. 12, 2MI :WI.
H. y" IIAMILTON. II. (1951) " ",'rips or 1I01'llial slag('!'> ill 1111' d"\l'IIlI'"II'1I1 of
1111' ('hiek J. fI.I"'/I},.. 1)9,.j.I) 92.
9. I'IA'I'T,.I. (195())Studi,'sOil Ihr prohl('111 or111'1'\'1' 1':1111'1'11. I.Trallsl'lalllalioll or1111' rOrl,lilld,
prilllordiu," 10 ('l'lopil' sill'S ill I\.lIIldy"lolllil../. 1':.1/,. /'",d.. 131,172 2ll2.
10. HOGI':IlS, W. M. 1I('lno\opil' spillal (,ol'd grufls ill 1lIIhry". Zoii/u)!y.
20, H7 219.
II. SI'I.:IlIlY, H. W. (1945) 1{('gUlllliYI' fal'lol's ill 1111' ordnly 1!I'0\\Ih or IIc'lIl'al cinliils. (;rou'lh
S.YIII[>., 10, 63 H7.
12. STHA7.NICKY, K. (1963) IlIvt'sligalioli or Ihc' fu('tilill or Ial'll'rolopil' spillal ('orll sl'gllll'lIls
ill IIII' ('nirk. AI'III Bio/. JlIITI)! .. SIIIII)I .. 5, !l(l.
13. SZI::KELY, G. (196:l) 1"1111('1 jOllal slll'c'iri(,jIy or spill"I c'ord "I'gllll'lll" ill 1111' "0111 rol or lilllb
1II0VCIII(lIls. ./. /';.\'[>. tl//Jr!,".. II, 111.
11. SZI::KELY, G., ./.(IW,2) ({pl'I('X "lid IlI'ha\'iolll' I'allt'rlls(,lil'ilcII frolll illlplalll-
('d slIpl'mlulll'n!l'y li.llbs ill 1111' (hjrk. .f. I;m/lryol. 1:.Ip. .H",/,/I.. 10, IWI:;I.
15. 1'. (19!lS) Nl'urol-(l'llt'sis ill 1Il1alysi!'> of d.V('lol'llll1I1. III \V1I.I.JIo:\(. B. II.. \'rEIS__ .
1'.1\. . IIAMUI'III;EIl, V. !\l'rvolls Sysll'llI. \,'\\ YII..I,.
D1 E FUNKTION IIETEHO'('IlI'ISCIIEH II i"I:I.. Io::\'\L\H1.. Jo:\TE
Ill':r i"11',<:I110:"
I1l'i lIiihllcTrlllhryolll'1I d('1' Stadil'1I I;' III (IIA\1II1 \(I:Io:n JlAMII.TON) \\1I1'11t'lI dir
op('l'nliv elllfl'I'1I1l'I\ Brnl'hilllsrgllll'lIlC' dl's IVl I,d1I11011'1'1llln's dun1I Ill'a('hiall'. Ihonlt'all' 1I11d
IlIlIIhosarralc Mcdllllnrrohrnhsdl/lillt' gll'il'llill'tig('l' :llId"I'I'r Jliilllll'!'I'lIIllryolll'lI ersllr.l. 111111'1'-
vatioll IIl1d FUllktioll drr dll\'('11 sol('III' Implalliall' WlIl'dl' lI:lI'h dl'llI AilS-
s('hliipfrll fiN lTiihnchl'n 1I111('rSllrhl.
Ar/II /li"l. /l1I1IJ!.. /-t. 1%:1
153
.... ill"s alHll'n'1I EIIII'l"y"S kiilllll'Il dit, ,'III f\'rlll {'II Brarhi .. lsr.gIllt'lIl I' d,'s
\\'irl"li,'n'" ill .il'd,'r 1I,'zi .. hlllll!: IllIll,,.valioll lIud Jo'lIl1ktioll tI"1" Jo'liigt'1 t'rwit'l;cIl l;i,'h iu
slllt-h"11 1"iill"11 al" uorlliai.
OII\"JI" all SI"II., d,'s ,'IIII'",.lIt .. 1I l\,.a('hiall .. i1s illlplulI\i,',.\., thoru('all' S,'glll"lIlt' di.,
1"ljig.. ( illll"1"\ i('rl"l1 IIl1d di"sl'lllt'lI sl'llIllI'rz"llIpfilldliC'h II un'lI, wa,. ihn' MIISrlllulllr voll-
kOIllUll'1I at ,.ophi",'h 1I11d ,'" "III II iC'''''' I,' "i"h "jill' 1\ Ilk ylos.' d,',. (;,,1(,11 kl'. 1);1' "illl,rhiiru,',. d,'r
illlplallli,'rl"l1 Hii,k"lIll1al"ks"l'/-.(11II'1I11' z"il!:l"11 wold ,'ill" d"lIlli,'h" IIYIIl'I'lrophil', di,' vortl,',.
!liiru"1" \I un'lI d:II.("I.(,'/1 at rophis('h.
1"(jig.. 1. dil' dllnh illlpialltil'rll' IlIlIdJO-sul'l"al,' S,'gllll'lIl .. iUIH'rvi.'rl WlIl"d"II,
J.'digliC'h illl St'ltllll""gl,I"lIk 1\I('iu"I"I' sll'ls lIIil dl'lI Sl'hrt'illH'wl'gllugell del' gll:iehs"iligell uul,'n'lI
(;li,'dIUUSSI'1I syul'hrolli"il'rll' .. 1H'r kl'iuI'de; fiil" di., sOllsl I ypisd\(' BI'\\I"
gllUl.(l'1I 111111 B,'I'II'x,', Tml;., gill allsg,'hilllt'ler lIud lIorlllul illul'l"vil'l"ll'r Fliigl'hllllsklllalllr wart'll
ill dl'lI 011111"/'('11 (;I,I"lIk"1I k,'i'll'rl.,; kool"llillicrlc BI'WI'gUlIg"1I lIa .. hweishar. Erkliirnllgsillii/-!:Iieh
k(';l"11 di""l'r 1I,'I'II11dl' 111'1"11,,11 diskulil'rl.
IKe.III':; toHA 11111': Y I 1I )lllJ <I>Y Ill{llltH1 ECI{11 x CI 11111 HO-
,\\():WOB1>IX CEI'MEHTOB
II I11,11 I.'IH"I' (LTa:1I111 I;' Hi 110 re/}.,q)'p?l'P.1' II rClMlI,'lJl1l1J/I)'} lIPCiUlClPIITl'JII.1I0
YHa:lclIlIl,ll' 1I.'IC'ICIlJ,IC CIIIIIIIIO\lO:II"OIll>IC ccnIClITI>' "J,IJIII :IilMClIlCIIJ" IIJIC':CIlJ,IMII, rpYilllJ,IMII.
IIJlJI II0>1l'1I1 I'II IlJ-1\ pCl'Tlllllll ,1.\1 I1 l'l'r,\ll'IIT'I.\I11 l'1I1111110ro M(1:II'(1, 0'1' HpyrJlx JapoJlJ,llIIei\
Ollllllal\OllCH"O 1l1l:lp<lCra, Il1Ic.'lc 1ll>lJlyll.'ll'IIlIH 11:1 )Iiillil I/CCJICilollaJIIICI. JIIIIICpIIRIUI)1 JI tllyll'<IUH/
l<pJ,I.'II'CIl, 11I1I1Cl'lll1iHlIl,1I111J,IX 0'1' ccrMClITOIl Clllllllloro MO;]ra.
11:lc"l'lloii ccr.\ICIiT ;qly,oro lllolllJlClIl,a .\llI)l<l'T :laMl'III1TI. Yila.llclHlJ,lii 1I.'IC'ICIIOii CCI'-
\1CIIT 111I IlCl:x lITIIIJIIICIIIIHX. B l'.'ly',allx lla:llllllla.'lacl. COIlCPIUCIIIIO 1I0p,\la:IIoII,1>1 JlIIlICP-
llallllH II IIIYIII\lUI>1.
XlI"n1 IICPIlIoI IICpCl'aiI\CIIIIJ,IX rIlY;lllJ,IX Cl'f \lCIITOIi l'a:lIll''I'IIJI)IJIIICI. II I\PIoIJIIo)IX, I\OTOpJ"C
CTClJIIi '1YIlCTIlIITC.'II,IIJ,I.\\11 II OTIIIIIIICIIIIII 110.lleiI, ,\11,1 1111 lJ,1 CTaJlJI COlll'PIIICIIIIO <lTPI)(11f14CCI<IIMII II
eycrillll,1 B il0I'Cil.'II,II0,\I l'l'I'0:'vl IlClIlCCTIlC 1Il'l'ccml<cIlIlIoIX ccrMCIITOIl lIafiJl/o
IlaJI<lCI, 1'IIII(:pTl'lllIIfIH, Torila I,al, II ;llllll'aTC.lII,IIII.\1 IIcllllC OllpCilCJICIIIIJ,IC IP":IIWI<l1
:U ICTpe It III Ill.
I'I'I>I:II,>IX, 1I1111l:IlIIJlpOllalllll>IX lJT rl:Tl:IHJTlJlIlI'ICCI<lIX 1I0)ICIIII'IIIO-I\PCCTllollJ"X ecr-
MellToll. lIaii.'lIoila:IIIl'I, y.\lCPCIIJlI>IC ;1111 I>ICCIIl IH IlJlc'!a, ClIllCIHIIClIlIlJ CIIIIXPlJllIlI.IC C IllIll)l<CIIII)IMII
1101' Toii i1,C noI' lJII J,I, 111I11I1(:1\1I XOIHIIIIO lla:lll1rroii II 1I0p",a:II.1I0 IIl1l1l'llIlllpOllatlllOii ",ycl<yml-
Typlol Ill: y;la.'lol'l. ynallollllTI, I\OOp;lIlpllll<llllll,IX :llllliI<Clllli\ IlPYl'JlX cyCTalloll, OiicY)I<ilacTcll
"pcIIIIlI.'larill'.\\I,lii alll\lI.'IO:la illICT<lJlhllJ,IX CYLTilllOIl II CII"XPOIIIIJ,IX illllDl<C/I/lii Il
IIJIC'IaX,
f<llI I/i"'. 1Il1"g. II. 1963