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Histogenesis of muscular tissue in amphibia I. Development of striated muscles from mesenchyma in urodeles

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This p p r i s the sequel of histologic iiivcstigations p u b
lished sirice 1929 (Katznelsoii '39, '31 and '32). Thc Amphibia
are L I S at~ present very g(mwilly as niatci*ialin cxperirriental
embryology, and it is from the urotlelan Amphibia that the
chief results in cieveloimcntttl mechanics havp loren obtained.
rlliese
?
investigations naturally lead to an attempt of influencing experimentally the course of histologic proccsses.
Expcrirneritj of this type require a previous detailed knowledge of iiorrrial histogenesis, altlioiigh we have a11 iiisufficient
acquaintance of the histogenesis of even some of the most
important tissues. A R regards the developmcnt of tissues in
higher vertebrates, we possess a certain amount of data ; but
our knowledge of the histogenesis of tlie tissires of lower
vertebrates is fragmentary. This is especially true as regards
the urocielan Arriphibia. Thus tlic study of histogenetic
processes in this animal order certairily rrierits consideration.
The large dimensions of the structiiral elciiierits in the L'rodela make them most favorwblc material for the elucidation
of a number of debatable histologic prohlcrns.
Much work has been done already 011 the histogenesis of
striated muscle-the problem of our present interest. It is
generally accepted, following F. SIaur-cI- ( '04),that the m p tome is the source of striated musculature. He writes in
I09
110
Z . 8. KATZNELGON
the chapter on the development of the muscular system in
0. IIertwig’s Handbuch (Bd. 3, T. 1, 1906): “Aus deli
Elernenten der Urwirbelwwduiig bildet sich die quergestreiftc
Nuskulatur des Rumpfes, S c h a n z e s und der Extremiliiten,
und nusserdem eiii grosser Teil der erribryorialeri Bindegeu-ebsanlagen” (p. 1). Since it is generally agreed that the
myotomes arise from cells disposed like epi tlielial elementsMaurer distinguishes striated niusculaturc as a n epitheliagenetic tissue and smooth musculature as a tissue arising
from nieseiichymal elements. “Man hat die Xkeletmuskulatur
der Wirbcltiere als epithelogerie Nuskulatui- bczeichnet
(0. Hertwig). Sie geht BUS der epithelialen Waiid der
Urwirluel hervor, im Gegensatz zu der meseiichpiatiiseii
Muslii~latnr,deren Elemente, die glatten &Inskelzellen, sic11 ails
einxelnen, im embryoiialen Bindegewcbe zerst rcut liegenden
Zelleii bilden” (p. 2). This ~ i i swritten 30 pears ago and
is still the predominant and generally accepted point of view.
G. Haggquist (’31), thc author of the section on muscular
tissue in W. v. I\lolleiidorfT’s IIandbucli d. mikr. Anatomie,
commences the chapter on the development of striated milscles as follows :
Die Skeletmuslielfasern werdcn aus dem dorsolateralen Teil
der Urscgmente d. sog. Muskelpla , entwickelt. TJiitersucht
man dereii Bau auf einem so zcitigcn Stadium, wie da das
zugehorigc Urscgment eberi beginnt sich abzugrenzen, findet
man deiisclben aus einer Sammluiig epithdnrti.q gelagerten,
polyponaleii Zellen zusammengesetzt. Diese Zcllen sind die
Rildungszellen der kluslielfascrn, die &lpohlastcri (p. 167,
italics mine).
Thus a strict distinction between the eyit2icliogenetie inusculature and smooth musculature of mesenchyrrial origin,
clearly delineated hy illaurer, may ex-en at present be considered as the prevalent point of view. In the special literature, at the lime w h i i 118ggyuist was writing his compendium,
are to be fouiid certain iridications that part of the striated
inusculature deve1ol)s independently of the myotomes, being
formed by means of metamorphosis and dil‘ferciitiation of
nicsenchymal elenleiits. Sucli data were ~)roducedf o r tlie
musclcls of the liead? extremities aiicl certain other organs.
As regar& tlie musculature of the extremities, \V. H. Lewis
(’02) ~ v a samong the first t o represent sacciiictly in l i i h paper
on the clevelopiiieiit of the a r m iii inaii the meseiichqrnal origin
of its muscles. In his analysis of the relation of the myotoniea
to the a r m bud, Lewis comes to the follomiiig coiiclusioii :
No distinct myotome burls take part in the formation of
this tissue (the tissue from m-liieb tlie mirscles, ligaments . . . .
of the a r m clewlop) (p. 153) . . . . ‘The arm bud is a t first
s ail cl closely pa clrrcl me seiichyma.
filled with :i liornog~~iieou
KO mi-ves or myotoine buds liare eiitered tlie arm, yet it eoiitains the tissue from \vhicli the miisciilar elements develop
(p. 181),
Among tlie recent litwatnre some not es1)eciall-y clearly
deliiiea tecl indications a s regards the mesrnchymal origin of
certain striated muscles a r e to be found in Tello’s paper
(’22) ; but it is to Victor ~ d i n i i d t(%) that \ve are iac1el)tetI
for the clear formulation of the new p i n t of view. The last
author published sonic 10 yca1.s ago ti prelimiiiary comniuiiication 011 the nieseiicliymal origin of the niusclrs of the estremities and of the tongue, aiid so1~1e117hatlater (’27) appeared
his \Toluminous paper 0x1 tlie saiiie problem. In this latter
work Schmidl clearly demonstrates that tlie 1iiu~c1csof the
tongue and extremities of manimalx are ill 110 may related
to the myotoriie musculature, Imt develol) directly f corn the
nchynia. The cmrse of the histogenesis of these n i p s enchymal imscles inclucecl liirri to abandon tlie conception
‘myoblast ’ iiitroclncecl by Qodlewdci, Jr. ( ’02).
AIeiiie obeii angefiihrteii U i i t e ~ s n c l i i x n ~ e i i - ~ ~ r i tV.
e~
Schmidt-liaben ergelwii, class die SI uskclfaseraiiln~eiii n den
Estwmitiiteii und in dcr Z u n ~ evon Siiiigetiureii sic11 aus drin
allgemeinen Illesencliymnetz differeneieren, class in den f riihpn
EiitT~~icBh~iiffsstadieii
die Extrcmit~tena1i1ttg.c.ausfiillt. Icli
sche lieiiie ‘ Uyoblasteii’ oder ‘XyozelZpn,’ d.11. voii clem
lIeseiicli~-miietngcsondcrtc, durcli ilirc Form nnd Struktiir
such untersclieicleiide Elmiente, die in c h i Rlcsencliymnctz
liogui u1~1
sich z u Jlnskelfnsern aushilcleii (p. 181).
112
2. S. KBTZNELSON
V. Schmidt ( '30) confirmed his investigations on iiiaiiimals
hy the studp of the histogenesis of striated muscles in a
rel~reseiitativeof a differelit class of vortel.mttes-tlie hippocampus. He demonstrated f o r the latter, the rncsenchymal
origin of the muscles of the pectinal fin, ~vhilethe course of
the histogenesis of striated muscle, i n general, proved t o be
identical with the motlc of tleveloprnrnt of this tissue in
mammals.
At present the notion that striated ini1wle tissue may
originate from mrseiichyma is already to he found in some
of the textbooks. Thus Petersen ('24, 1). 2 x 1 ) states that
striated muscles may originate from myotomes, as well as
f r o m mesenchyna. Strong and Elwyn in Bailev's Textbook
('25) mcntion the cephalic muscles aiid those of the extrcrnities a s possibly originating froin mcr;enchyma ; as rcgarcls
the extremities, l i o w ~ ~these
~ r , authors surmise that the me$enrhyma migrates from myotomes. A. A. Zawvursin ( '%,
p. 298) is of a similar opinion ; he nimtioiis Schmidt 'Y paper
but is somewhat skeptical as to the possibility of a differentiation of muscles directly from the mescnchpnia. ,J. SdiaIler
('32, p. 301) indicates a number of muscles for wliich a
mesenchpnial origin is possiblr. It is, hotrLrvcr, charactcristic ihat all the tpxt-books Iierc mentioned describe orilp the
deveIopiiierit of striated iiiuseles from myotomes through a
' my obla st ic ' o ri gi11.
Some time ago, the question as t o whether the striated
rriiiscle fiber is of uriieellular o r mnlticellnlar origin was rnucl]
discussed. This problem is still unsolved (for detailed list
of literature see k'ranx ('15) aiid Hiiggquist ('31) : in the
latter compciidium a nuiuber of papers have not beer1 cited).
Some authors still favor the theory of unicellular origin of
the striated iiiusele fibers (IIiiggquist, '31 ; Rchaffer, ' 3 3 ) .
Others are of the opinion that in different animals there are
present different types of development of striated muscles.
Frariz ( '15) considers the musculat in-c of invcrtehrates to
be of multicelliilar origin, while h e ascribes a nniccllular origin
to this tissue in vertebrates. A unic.ellulau spucytial develop-
ment has been described for mammals by Uocllewski, Jr.,
( '02) aiid Mlodowska ( '08). The paper of the first 01 these
two authors is considered as a 'classical' work on histogenesis
of striated muscles arid a s such it is cited in nearly all the
text-books. However, even the authors that stand for syiicytial
ctcvelopmeiit of striated muscles in mammals as described by
(:odlewski a r c of aiiother point of x-iew as regards Aiiiphibia.
They consider that in this latter class tlw striated musculature develops from a imicellular type ( TT. Enbashkiii, '31,
pp. 254-255; A. Zitwarsill, '33, 11.298).
Aniong the older authors, R. Remak (184.5),Weissmauii
(1861) and Kollilicr (1861) have investigated the problem
of histogenesis of striated rnasc1c.s in Ainphhia. lister,
l<yclcsli>-mer ( '04) iiiatle a study of the dcvelopmc~ntof muscles in Neczturus. The most detailrd work, ho~vever,belongs
to Franz ( 'I.?), w1io studied the developnient in the urodclari
Xmphibia (Triton). Recently, two papers on histogenesis of
striated muscles i n Amphibia (frog) have heen published by
Haggquist ( '20). Eg'cleshyrner, Fraiiz, besides Haggquist
studied only the myotomal dorsal mnsculatare. In none of
these papers is mention made of the meseneliymal development of striated muscles in Amphihia. The authors here
named are of the opinion that the niuscular fiber originates
from niyoblasts, myotome cells lying like epithelial elements.
Each of these cells retains its separate character and forms
a striated muscle fiber.
From this cursory review of the literature two points may
be inferred: 1) The possibility of a rnesencliymal origin of
striated muscles is still coiisiclered by most authors as
problematic. 2 ) In relation to , h p h i h i a there a r e no inclicatioiis of the occurrence of such mode of histogenesis of striated
muscle and f o r them the derclopmcnt according l o the unicellular type is gerierally accepted.
In this paper T propose: 1) To confirm the fact of a mesenchymal origin of striated muscle in vertebrates. 2 ) To
show that over and above the unicellular mode of dcvelopmcnt
of striated muscle fibers, described by the authors that have
114
2. S. IlATZNRLSON
studied the myotonial histogenesis in Amphibia, there exists
another mode of development of striated musculature.
MATERIALS A N D hI1ETHOl)X
As in my previous studies I used f o r material the larva
of the Ural salamander, Salamaridrella keyserlingi Dpb. The
fixation of the material was accomplished either with Zeriker 's
fluid o r a modification of this fluid according to Helly
(Zeiiker-formol). I n either case the fixation was prolonged
for 24 hours. Both fixation fluids gave particularly good
results when used for such developmental stages in which
there was present but a small amount or absence of yolk.
The fixation of the larve was accomplished in toto, tlieii they
were washed, hardened in alcohols and embedded in paraffin.
F o r an intermedium we used mostly clove oil,l less often
xylol. Series of sections cut i n different planes tlirough entire
larvae were prepared, the sections being of 6 p thickness.
The preparations were stained exehisively i n Heidenhain's
iron haematoxylin, using occasionally as a plasma stain
Congo red or picrofuchsin. The figures wew delineated
personally by nieans of a camera lncida, Zeiss.
OBSERVATIONS
According to my observations, formation of striatecl miiscles
from inesenchyma in Ampliibia takes place in the development
of the muscles of the head, gills, the ventral ixut of the
abdomirial u d l , and those of the extremities. The COL~PSC of
the primaq- formation of the tissue is, iii the maiii, similar
for all the muscles that have been here enumerated; it snffices, therefore, t o give a general description of the meseiichyrnal hist ogencsis of the striated muscle. This description is to be based principallyy on the data of the development
of the muscles appertaining to the extreinities. The differentiation of meseiichynia that leads t o the formation of striated
For the description of thc rapid method of cmbcdding in paraffin with oil
of clove as an intermcdiuru see my former papeis (Katenelson, '31, '3.5).
a A short prcliininarj communication on the iesults of this work, but without
any illustrations, has been published in the C. r. Acad. Scient. 1-RSS, 1934.
HISTOGENESIS OF MUSCLE I N A N P H I R I A
115
muscles does not take place simultaiieously in all the above
mriitioncd points. Thus it is that, for instance, in the extreniities, the mesenchymal miiscles appear after the cartilagenous
Fig. 1 Mesenchgma of the cephalic end of an axolotl larva (cighth day of
derelopment) . Fix. Fol's method. Heidenhaill '9 haematoxylin. Oe. 2, Obj. imrii.
18C Rciehcrt.
skeleton has bceii formed. Namely, at the time when part
of the meseiicliyrnal primordium of the extremities is already
differentiated. T t appears that the mesenchymal iiiuscles
tlevelop Iatcr than thc muscles of myotonial origin. 111 larvae
116
Z . S. IihTZJSELSON
that show but tlic first signs of a museular differentiation of
the meseiichynia the myotornal muscles are already full:r
formed.
The structure of tlie mcsenchyma in the early dwelopmental
stages of thc urodelan Amphibia has lieen descrihcd in one
Fig. 2 Bud of the anterior catrctrlity of a Salaiuunclrella larva (sixtccllth day
of drvc~lopmrwt, a h o u t 10 i m i . long) . Frontal section. Zenker. Heidenhaiii ‘s
haernxtosylin
Congo red. Photo.
+
of my 1)rcvious papers (Katznelson, ’22, pi). 950-951) from
wliich I have borrowed figure 1, showing tlie character of
this tissue in Axolotl (the same is also true f o r Saltmiandrella) . 3 The loose mesenrhymd tissue rcscmbles an opeiiwork syncytial network in the l’lasmatic nodides of which the
nuclei are to bc seen. Tliese muscular plasmatic territories,
This description coincides rritll the dcsriiption given b~ F. P. Mall ( ’ 0 3 ) f o r
the meseiichyina of the tadpoles of the frog.
HZSTOGENESIS OF M U S C L E I N A A l l ' l l l B l A
117
it secms, possess ihe faculty of scgregnting from the entire
syvncytial network ;ml chniige i d o Iticscncliq-ma1 cells : 011 {lie
otlicr hand, nndci. certain circurristanres such cells may agiiill
take part in the composition of the general syncytial networl;.
In other places the mesenchyma accumnlates in cl~iiscr
masses ; in this case only ihc protoplasm around the nuclei
(endoplasm) mag- become visihle, while the peripheral 1)mt
of thc, ~ ) r ool)lasrn
l
(ectoplasm) acquircs the character o f the
Fig. 3 Scctioc forming part of n series through a Salarnniidrella l a ~ r a(thirtysecond day o f development, lo.., Inm. long). Zenkrr. Heidenl~ain's liar~mntosrlin
Congo red. Primary rlifl'ereiii iatioii of the nicsciichyinw in the region
of the anterior extremity. Acemnulation :id p o h r orimt:i,tion o f nuelpi. Oc.
X 15, O h j . 40 Zeiss. X GOO.
+
intermediary substance and its proccsses disalqwwr. f3uch
is the appearance of the mesenchyma i n the hncl of the extremity (fig. 2).
The first indication of 1lie begiiiiiitig of differentiation of
the mese~ichyiiia toward formation of striatcd muscles consists in the appearance of local accumulations. The mcseiichymal cells (the nucleated territories of the syiicyt ial
mesenchymul network), dispersed in the beginning come to
lie more or less parallel, while the nuclei dispose themselves
Compare HtudniCka ( '33).
118
7;.
S. KATZNELSON
in such a manner that their longitudinal axes are similarly
orientated (fig. 3 ) .
Fig. 4 Taken from the bud of ihe anterior cxlreiuity of a Snlaniniidrella larva
of 23 mm. Zmker-Formol. IIeidcrlllain 's haematoxylin. Longitudinal (lower
right) aiid tranmcrse (upper l d t ) sections through a muscle himdlo iu its
enrly stages of developmcnt. Oe. x 10, Ob-j. imm. 90 Zriss. x YOU.
Thc study of the preparations shows that in some cascs
in such groups of similarly orientated nuclei no figures of
division are to be found, while in other cases, on the contrarv,
8 great number of mitoscs ( u p to seven in oiic optical field)
inay be observed (iig. 4). The initoscs in sucli amassments
of meseiichyma arise simiiltaneously in sort of cxplosioiis.
T7cr37 ofteii the lxotoplasm around the iiuclci segregates and
distiiictly contoured rouiicled cells come t o be formed. I n tlic
cells not undergoing divisions the protoplasm is visible only
at the 1)oles of the longitiildinal nucleai- axes ( endoplasm),
occasionally nearly the entire protoplasm may assume an
ectoplasmatic character, i t heoornes entirely imperceptible
in some places and the nuclei appear to be bare.
Fig. 5. ‘I’mo iiiielri from
mosele bundle in the early stages of dcvclop~nent
frorii an anterior extremitj- at the stage of t h e appearance of iiiyofibrillne. Zcnker
Foiinol.
IIeidcnhaiii ’s Iiarmatoxylin Congo red. 1, ordinary mrsenchgmai
nuelcus. 2, vesicular nuc1en;s. Oc. X 15, Ohj. imm. 90 Zriss. X 1350.
+
Thus iii tliis l r i n i a r y stage of differentiation we distinguish
t w o riioiiieiits : condensation of the iiuclci aiid llicir polar
orientation. The condensation alone denotes local differentiation of riiesenchyma ; the direction in which this differciitiatioii takes place remains as yet unlinown ; slill, the
siiiiilarity in the orientation of the nuclei may he considered
as a first sign of the fact that the differentiation of the meseiichq-ma tends toward Ihc formation of muscles.
Already at this early stage the presence of degenerating
musclcs is to be iioted. Heidenhain’s liacmatosylin stains
such iiuclc4 illtensely ; it s e e m tliut the degeneration of p a r t
120
8. 8. KATZNELSON
of the nuclei -must be considered as a procws that inevitably
accompanies the embryonal differentiation of
The following stage of dcvcloprricrit of striated muscles from
meseiichyna is characterized by the appearance of myofibrillae. Sollie of the mesenchymal nuclei in this stage
Fig. 6 L on g i t u d i n a l section through a inuscle buiidle from the proxima1 region
of a n anterior extremity (Sulamandrelln. larva, 2 3 1mn. long). Zenkcr-F'orinol.
11eideiihnin's haematoxylin
picrofuchsin. Oc. X 10, 0b.j. iirim. 90 Zeiss. X 900.
+
undergo characteristic changes. Their chromatin disappears :
there reniaiii in the riuclci only a rounded karyosome (nucleolus) distinctly visible against the light, transparent background of such nucleus. This sort of nuclei a r e easilF
distinguished from the rest of the mesenchpmal nuclei j these
latter a r e rich in chromatin which takes the form of small
Compare b h i s t ( 'M), Kallius ( '32), Katznelson ( ' 3 2 ) and Muhlniann ( ' 3 2 ) .
HTSTOGEA-ESTS O F MTTSCLE I N A M P H T B J A
121
masses of granules (fig. 5 ) .6 One of the poles oS tlie 'vesicular
nuclei' often bccoiiics soiiiewliat elongated and it is exactly
at this point that tlie diffeiwitiation of tlie myofibrillae sets
in (fig. 6 ) . The latter appear as bundles of fine undulatory
filarnerils traversed by stronger iiiterisely staining fibrillae.
All thc fibrillae run in thc direction that coincides with the
axis passing through the nuclear pole.
The older authors (f.i. Wagener, 1861) were of the opinion
that primarily the fihrillae arise in f o r m of homogeiieous filaments. Later authors described a granular forestage in the
developnieiit of the fibrilla e (Godlcwski, '02 ; Jllodowska, '08)
and saw the homogeneous filsvillac arise from the fusioii of
these granules. Tlie striatioil of the fil)rillae makes its appearance later and repreheiits the fiirther differentiation of the
formerly homogeneous fiber. 31. llcideiihaiii ( 1899, '11) also
shares this point of view. Eecently, however, IIiiggquist
('20, ' 3 3 ) has dccidedl>- ol~poscdthis opiiiion. In a special
paper on the dcveloprncnt of myofibrillae iii f r o g ('20) lie
most ernphatical ly asserts that the striation of the myofibrillae
is a primary plieriomciiori and denies the existence of the
homogeneous fibrillw stage. My ow11 observations do imt
tally with Haggquist 's data. I n tlie developmenttd stages
described I have always observed in my preparations the
appeararice of Iioniogeiieous fibrillae that oiily later on transform themselves into striated fihrillae.? At thc cnds of these
homogeneons fibrillae a sort of bunch-like split tirig is observed
* The modifications of the nuclear structure duiing lrlrogenesis in Airiphibi;t
haTe been specinllq- studied by F:~cleshpier ( ' @ 4 ) , but he never saw nuclei
resembling the 'vesicular' nuclei I hnxe deseiibed. On tbe other hand, ir. Srhmirlt
( '27), has obserx ed i n maiiiinals in tlie dcrclopirient of stiiated muscles fronl
the mcseiich~nia.some similar nuclei.
'LIlggquist is of the opinion t h a t the reason wliy striation has n o t been
observed by other authors lies in tlieii ha>ing used an inadeqnete stain (Heidenhain 's nictliod) and recoinrnends staining with iron haeiiiatosylin nccordlng t o
the method of Hnnaen. Jt reniains, home\er, unex~~laiiied
why i t is that in tt
lmparntinn treated with Heideiibain 's hneniato\ylln u f two fibrillae lying in
close vicinity to one another t h e stain is capable of showing striation in one of
thein and not in the other. One ran say t l ~ tth e only possible caplanation of this
fact is that in one of the GlJrillac the basis f o r the phpico-chemical diEercnt1;t
tion of the discs is already present, wide in the other such differentiation llits
not taken place.
THE A64TO\LIP*L RFCOIID, VOL. 61, NO. 1
122
2. S. IiATZNELSON
that does not occur in the iibrillac that display striNt’1011.
The homogeneous fibrillae stain more sharply than those that
a r e striated. The fibrillae end in tlie vicinity of the nuclei.
but sometimes they are bent out of their course by the latter.
Fig. 7 Transvcisc wetion through a niusclc bundle in its early stage of drvelopmcwt froin thc ventral region of tlic abdominal wall (Selamandrrlla larva,
2 3 mm. long). Zenkcr Forinol. IIcidciiliaiii’s haematoxylin. Oc. x 10, Obj. inmi.
YO Zriss. X 900.
Figure 6, illustratiiig tlie facts here descrihed, shows it
longitudinal section of it muscle bundle in its earliest stages
of dcvelopment. Figure 7 r e l m s e n t s a cross section of such
a buridlc ai the same stage of development. Here the ‘vcsicular nuclci’ with their karyosomes represent 811 outstanding
feature of tlie microscopical picture; they lie strewn among
the ordinary nuclei of the rnesenchyma. Between the two
HISTOBEPiESH OF M U S C L E I N AMPIIIUIA
123
types of nuclei, transitional forins a r e plainly visible. Tlic
protoplasm surrounding single iiuclei as me11 as groups of
nuclei appears in the form of irregularly contoured districts,
these contain deeply staining points that represent cross
scctions of newly formed fibidlae lying together in thin or
thick bundles. Occa sioiially single fibrillae may be discerned.
The next (third) stage is characterized by t h e forming of
muscle fibers (Primitivhiindel of German authors). They
originate through longitudinal splitting of the myofihrillae.
This has hccn noted by a number of authors, and RI. TTeidenhaiii (1894, ’11)has made a particularly detailed study of it.
Fig. 8 I’riinnry muscle bundle f r o m an extremity of a Salaroandrelln larva
(fifty seventh day of development). Zenker. Heidenhain’s haeinatoxplin. O r .
x 15, Obj. imm. 90 Zeiss. X 1 3 5 0 .
The splitting begiiis in the middle of the fibrillae, as it has
been in latter years described by Ilaggquist (’20). Recause
of the nature of the splitting tlic buridle of dewlopiiig fibrils
assumes a spindle-form. Contiguous to such a spindle-form
bundle of fibrils and laterally to it lie the nuclei surrounded
by a certain amount of uricliffcrentiatcd protoplasin (sumoplasm) (fig. 8). The nuclei here possess again their ortliiiary
character. I have never found ‘vesicular nuclei’ at this stage.
Tn opposition t o IIiiggquist (’a()), who has observed a sarcolcinma in the earliest stages of myogenesis, I could not discern
a sarcoleninia even in such stages as arc represented in this
figure. The study of later stages did iiot interest nie, hiit
it is evident that the sarcoleniriia appears much later, possibly
in connection with the differentiation of fibrous con11
tissue. At least in none of my preparations have I ever met
124
2. S, HATZNELSON
anything resenilding Iliiggyuist’s double contoured (sic !)
primary sarcolemnia (‘ priniiii-es 8arkoleinm’).
ils regards striation, the figures here given show clearly
that in the beginning it comes o d y to the formation of the
P i g . 9 Part of a section through a gill arch (Bu1:iiimidrella larva, thirty
second day of dcrelopmeat, length 10..5 nim.). Zmkrr. IIeidenhain’s lracrria
toxgliii
Coiigo red. Oc. x 10, Obj. imm. 90 Zeiss. X 900.
+
discs Q and 111, while in the stages stuclied the lines Z and h l
have never come to rny observation.
Certain special details of the processes described iriay be
meiitioned here in respect of the developnient of mesrnchymwl
muscles in the gills (figs. 9 and 10). There the fine niuscular
layer originates subepitheliallp. The developing muscle bun-
HISTOGEXEGIS O F M U S C L E I N A i U P l l I U l b
3 25
clle very soon takes on the character of a band along which
the nuclei come to lie occasionally ovcirlapping. one another.
The nuclei show the charactcristics of ‘vesicular iiuclci,’ as
described above. Here also hoiiiogencous Gt)rillac are p1-iiiiarily t o be observed, the striation setting in as a secondnr.y
phenomenon. In the gills long plasmatic rods conic t o view
along .cvhich the nuclei a r e s f r ~ w n but
,
the pro1oplasm shows
as yet 110 trace of fibrillar structurc (fig. 9). T n othcr places
Fig. 10 Part of a section throiigh a gill tire11 of R Salainandrclla laria, about
10 mm. long. Zciiker. IIcidcnhain ’3 haematoxpliii
Congo red. Oc. X 10,
Obj. irrim. 90 Zeiss. X 900.
+
I have never observed mch rods in stages 1)recediiig the differentiation of the fibrils, and the protoplasm is rather of
a spongy character.
The primary development of meseiichymal striated inusclcs
depicted hcrc diflers considerahip from the histogenesis of
the striated musculature as tlesczribcd hy the authors wlio
have previously studied this process in Ainphilnia (Franz,
’15; €I%ggqiiist, ’%O). Not wisliiiig t o untkrtake here a detailed examination of the question, 1 sliall mention only the
126
Z. S. KATZNELSON
principal poiiits of diffcrencc. A11 the authors mlio h a w
described the development of the striated muscles have considered as a starting point the formation known as ‘myoblast.‘
According to Godlewslii, the coiiception ‘ myoblast ’ or muscle
ig
forming cell (‘muskelbildende Zelle ’) has tlic f o l l ~ u ~ i imeaning : “die embryoiialc Eleinente, welche sich iiur meriig v o ~ i
den iibrigen embryoiden Zellen unterscheiden uiid welche
cliircli wcilere Differenzierung sich zu Aluskelfaserii heranbilden li6iineii” (’02, p. 114). Victor Schmidt (’27) indicates,
and his opiiiioii seems t o me pcrfectly wcll founded, that these
terms are not t o be applicd t o the mcserichymal development
of muscles because in this case he saw neither ‘myoblast’
nor ‘muscle cell.’ From the very beginning of meseiicliyninl
differentiation we find a spongy mass of protoplasm (or a
protoplasmic band as is the case in gills) in which it is in?
possible t o distinguish any territories o r segregated elements
t o which the term ‘illyoblast’ might be applied.
The authors who have described the histogenesis of striated
nmscles in Amphibia (Franz, ’15; Haggquist, ’20) consider
the muscle iibers t o be of unicellular origin. Leaving aside
the question in horn- far this point of view is correct in resl)ect
of the dorsal myotomal musculature, it is to be noted that
f o r mesenchymal muscles the unicellular theory is most decidedly unacceptable. Franz (’15) is of the opinion that the
niultinucleated structure of the muscle fiber is the result of
amitosis. A number of authors mention this mode of nuclear.
division-in particular V. Rchniidt ( ’27, p. 144), ~7hodescribes
it for ineseiichynial muscles. Personally I have never observed
amitosis, this may possiblj- take place later, hut in the stagrs
that I have stuclied the division of the nuclei in tlic muscular
bundle that is being formed takes place exclusively by means
of karyokinesis.8
’ Nicolaides (1883) has obseivcd kargokinesis only i n the earliest htagcs of
development o f the muscles. Godlcwski (’02) is of tlir opinion that mitoses
orciir also in the later stages and it acems that Kiilliker (1889) inrliiies toivard
the same point of view. Hhggqnist ( ’20) thinks that i n early stages the mitoses
are strewn along tli? whole fiber that I S being formed aiid latcr in the central
p a r t amitoses arise while thc ends of filJcrs may contain mitoses.
HISTOGENESIS OF' MIJRCLE IK AMPHIBIA
127
The generally accepted point of view (introductory p a r t )
that the earliest stages of derelopmeiit of striated muscles
in Amphibia differ from the genesis of this tissue i n mammals
is refuted by the fact that there is a certain rescmblurice
between the data conceriiirig tlie rnyotomal niusculature of
inarrinials as giveii by Godlewslii ( '02) and, according to my
own observations, those of dmphibia. Rut quite particularly
to be noted is that the histogenesis 1 have described of the
muscles in Amphibia, in its 1,rincipal features, coincides with
the observations of Victor Schmidt on the development of
the muscles of the tongue and the extremities in mammals.
By comparing the ohservatioiis of this author with my own,
one is led to the cordusion that i n these two classes the early
developnient, leastways of the mesenchyrrial rniisc1es, occurs
i n a n identical manlier. Considering the farther resemblance
of thcsc processes to similar processes that take ldace in fish
(V. Schmidt, '30) the supposition that in all Vertebrata ilic
striatrd rniiscle develop in an identical manlier becomes quite
plausible. In direct opposition to these coiiclusioiis are the
clcscriptioiis of the histogenesis of the mpotonial muscles in
A4inphihia giveii by Franz ( '15). Considering the data obtained as a result of these irivestigatioris of the early stages
of development of striated muscles from mesenchymtz, a
supplementary investigation of histogenesis of the myotornal,
dorsal musculature would not be amiss. I sliould say that
though, up to tlic present day, the data that a r e to bc found
i n llie literature a r e of the most contradictory kind, still
there comcs t o mind the supposition that in its general outliiic
the myogenesis must be the same in all vertebrates. Rerent
researches show that there is a certain siniilarity between
tlie early development of the rriyotomal and that of the ruesclichymal striated rnnsculature in mammals. Thus, Earl 0.
Butcher ( '33), while studying tlie development of striatecl
muscles and tendons i n the mpotomal iiiasculaturc o i the rat,
remarks upon tlie parallel between h i s own obsei-vatioiis
and the process of meseiichyirial myogenesis as described by
V. Sclimidl ( ' 2 7 ) . These data prove anew the necessity of
a reirivestigation also for Amphibia of the development of
the inyotonial muscles.
128
2. S. KL4TZ-UELSOPIT
I t lias been irieiilioiied in the iiitrocluctory part that Yanrer
( '04, '15) made a sharp distiiictiori between the epitlicliogenetic striated musculature and the meseiichymal plain milscle tissue. But, physiology tcachcs us facts that do iiot tally
with so sharp a clistiiictioii betwccri tlicse two forms of
muscular tissue. Carey ( '31) uffirriis lliat plain muscle may
experimentally be transformed into striated muscle tissue.
The study of the mesenchymal histogenesis of striated niusc~ilature bridges over tlic cliflerciicc existing betweeii llic latt CI'
and plain i~iuscle tissue. A s both these forms of muscle
tissue may be derived from meseiiclipia, it is to be supposed
that their histogenesis must have certain points in common.
I n that respect it may be apposite to make mentioil of the
coiisiderable rcseinblaiice observed by Victor Schmidt, as well
as by me, betweeii tlie liis togciiesis of mcsenchyrnal striated
muscles and plaiii muscles (viz., YkcGill, '08).
Tliis work treats of only a few 01 llie problems coiinecteci
Tvith myogenesis. It follows iiainrally that next must COIUC:
researches on the histogenesis of iiiyotoriial muscles, special
iiivestigatioiis by means of cptological mcthods, etc. Since
i n this paper I have limited my studies to the elucitlatioii ol'
tlie problem of ilie origin of striated musculature arid to the
comparing of the general course of rueseiichymal mq-ogeiiesis
in Amphibia with that of tlie other vertebrates, T intend to
take up this theme in my folloming papers a n d to malie n
particular study of the probleins resnl tiiig from the present
irivestigatioris that a r e to be considered as the first part of
a larger work.
SUMMABY
1. Part of the striated musculature of vertebrates is derived b~ means of local clifferentiation from the mesenchyma.
2. Of meseiichymal origin in Amphibia a r e the muscles of
the extremities, the gills, those of the ventral part of the
abdominal wall, aiid the muscles of the head.
3. I n the development of the mesenchymal s t r i a i d muscles
lliree stages may be distiriguisliccl: 1) the stage of condeiisation and orientwtion of the mesencliyinal nuclei ; 2 ) the stage
of the apl-wmmce of myofibrillae that stands in connection
HISTOGENESIS OF MUSCLE I N AMPHIRIA
129
with an alteration of the structure of part of the mesenchymal
nuclei (vesicular nuclei) ; 3) the stage of formation of muscular fibrils. The course of mesenchymal histogenesis of striated
musculature is evidently, in its general outline, identical in
all vertebrates.
4. The data concerning the development of myotomal musculature in Vertebrates requires to be investigated anew in
order to elucidate the typical ontlines of the histogenesis of
striated muscles, characteristic for the entire group of vertebrate animals.
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