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Growth of transplanted tissues derived from normal and overripe frog's eggs.

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OKOTTTH O F TRANSPLANTED TISSUES DEIlIVED
FROM SORMAL BND OVERRIPE
FROG'S ICGGS
J O H N E. ALLISON'
Depuj t m e n t of Zoology, The State Unzvers~tyo f Zowu, Iowa CLty, Iowa
FOURTEEN FIGURES
INTRODUCTION
Among manifestations of the adverse effect of retention
in the uteri on unfertilized eggs of a female frog beyond the
time when they would normally be laid are the impairment
of their mechanism of protection against polyspermy and
their inability to develop into normal embryos following
fertilization. The number of days that they must be retained
to become thus affected (overripe) decreases with rising
temperature.
The literature concerning the effects of overripeness on
amphibian eggs has been reviewed extensively by Witschi
( '52).
Belogolowy ( '18), Diirkcn ( 'as),Lehmann ( 'SO), Spemann
( '42) and Waechter ( '50) made homoplastic, heteroplastic
and senoplastic implants of normal embryonic tissue of
selected aiiamniote types into various regions of the body
of older hosts. The implants were either pieces of prospective
ectoderni or whole embryos in the blastula o r early gastrula
stage. Some parts of such implants become incorporated
into the normal tissues of the host, whereas other parts form
accessory structures such as muscle, nervous tissue and noioThis investigation was supported by n research grant from the National
Scieiice Foundation ; grant adniiiiisterd by Prof, Emil Witschi.
2 Preseiit
address: Anatomy Department, St. Louis University School of
Medicine, St. Louis 4, Missouri.
561
562
J O H N E. ALLISON
chord. k'requently, infiltrating iieoplasnis develop from some
of the cells of the implants. I n many instances, these destroy
sonic of the normal host tissue aiid in sonie cases cause the
host to die.
Rierich ( '%), Rautzniann ('29) and Iiusche ( '29) performed experinients similar to those outlined above. They
conclude that implants of normal tissue are not predisposed
to develop into malignant growths.
Diirken suggests the possibility that implants from overripe eggs have a particular ability to develop into malignant
neoplasms. liehmann believes that the lack of oxygen and
the excess of carbon dioxide in the environment during the
period that his implants were becoming established map have
altered t h t m in such a iiianiiei- as to cause some to become
inalignan t .
Witsclii ( ' 3 0 ) implanted p a r t of an abiiormal embryo from
an overripe egg into a normal larva. At metamorphosis, this
host had carried the implant for 22 days. The infiltrated
reticulum from this frog, together with parts of the organs
to which it was attached, was transplanted t o the body cavity
of an adult frog. A t the end of 62 days, outgrowths from the
tumor which it formed penetrated the host tissues, folloming
the reticulum. The tumor caused partial destruction of the
urinary bladder and produced metastases in the liver and
intestine of the host. JYitschi ( '52) suggests that inflammatory
reactions niay have played a role in producing certain aspects of tlicse results.
A characteristic teratoma appearing on some embryos from
overripe eggs is a n ectodermal growth resembling a papilloma.
Rriggs and Berrill ('41) grafted these to various regions of
the body of 11ormal tadpoles. Of 218 cases, only three developed papillomatous growths wliich appeared to be comof host ectodci-m; the hosts exhibited a retarded
development, characteristic of animals from overripe cultures.
The present work compares the developmental potencies
of embryonic tissues which are derived from normal aiid from
overripe eggs.
G R O W T H O F ABNORMAL FROG TRANSPLANTS
563
MATERIALS A N D METHODS
Female frogs (Rana pipiens) were kept at 4" t o 7" C. prior
to being used. On the day they were to be stimulated to
ovulate, their environmental temperature was gradually increased to 20" C. These females were injected with from
two to 5 frog pituitaries each. Usually a few eggs could be
stripped from the uteri on the day following injection; they
were artificially fertilized. Animals that developed from such
eggs mere used later f o r normal implant material or as
hosts. The remaining eggs were stripped and fertilized on
following days. Most of the overripe eggs were kept in the
females 4 days or longer.
I n one series of experiments, homoplastic and heteroplastic
implants mere made either into the body cavity or into the
orbit. Larval hosts were Rana pipiens and Rana catesbeiana
from stages 25 to 30 (Shumway, '40 and Witschi, '49 and '51).
Adult hosts, Rana pipiens and Rana palustris, were also
employed, and implanted with whole embryos or parts of
embryos of Rana pipiens. These implants ranged in age
from late blastula t o stage 21. Control implants were from
eggs laids the first day after the female had been injected
with pituitaries. Two animals received implants of embryos
from eggs obtained two days after hypophyseal injections.
The remainder of the implants into experimental hosts were
from eggs obtained 4 to 5 days after the female was injected.
Most of the implants were undisturbed until the host had
reached stage 30 or beyond. At this time, the implant and
some of the surrounding tissue was removed. Some of these
were fixed for histologic examination. Others were separated
as completely as possible from the host tissue and were then
reimplanted into a new host.
I n many instances the implant had increased to such a
size that only a part of it was reimplanted. The remaining
implant tissue and the surrounding host tissue was preserved
f o r sectioning. I n some, reimplantation was repeated until
the original implant had been carried through as many as
three hosts, None of the hosts carrying implants from normal
564
J O H N E. ALLISON
eggs \\-ere killed earlier than 1 7 days after inlplantatioii. Of
all the hosts which survived, 129 received implants of overripe illaterial and j 3 rcc>eived normal implants. Tables 1
and 2 indicate the iiurnber and kind of hosts used and the
sequence f ollowecl in niaking reiniplants.
HOSTS FOB PRII1,kRY
I3IPL9STS
HOSTS F O R S I C O N D A R T
IMPLANTS
( 27 R. pipieiix larvae69 R. pipieris larvac -
1
1 7 R. catesbeixna
la1va _ _
4 K. pipiriis :idtilt\
1 R. pipiciis adult
1 R. palustris adult
2 R. pipieris larvae __
IlOSTS FOR T I R T I 9 R I
IIIPLA8TS
'
'1
4 R. pipiens laivxc~
1 R. pipiens adult
1 R. pnlustris adult
2 R. l~aliibtiis:rdults
TABLE 2
, ~ i t ? i i1
b' ~n i i t l 7iint7
of Iionls recercrng implants of t m s i i f delired f r o m rnib?'!/os
dcveloped f r o m normal eggs
HOSTS FOR S E C O N U A K T
IMPLANTS
I[OST* F O R P R I M A R T
I.\IPIITTS
38 R. 1)ipi(ws larvae --
I
10 R. pipiens larvaea R. pwlustris adults
HOSTS FOR TERTIARY
IMPLANTS
2 R. pipiens larvae
1 R. palustris adult
In a second series of experiments, epithclioma-like growths
from overripe embryos (stage 17, fig. I), were transplanted
to the prospective flank region of normal hosts (stage 1 7 ) ,
fig. 1). Figure 2 represents a section through a typical donor.
A patch of ectodcrm about O.imm square was removed from
each no~inalhost. A piece of the abnormal growth from an
abnorinal donor was then placed over the ectoderm-free
area. This tissue from the donor included ectoderm and
lateral niesoderni, and was large enough to overlap the
opei*ated region of the host on all sides.
G R O W T H O F A B K O R ~ V A L FROG TRASSPLABTS
565
Control aiiiiiials received similar grafts of noimial ectodermal and mesodermal tissue.
Of the epithelionia-like series, 5 experimental aniiiials were
killed on the second day after the operation; 22 experiniental
aiiinials and 11 control animals were killed at the end of 24
days at which time most of them were a t stage 25; arid the
last 20 experimental animals and 13 control aninials were
preserved after having carried the transplant 51 days. Thew
were a t stages 26 and 27 when killed.
RESULTS
Iiitpluiits k t o lartwl hosts
Geizeral obsclrrcitions. In 16 of the implantation cases
studied, no foreign tissue could be found in the host at the
time of preservation. Three animals died. Eleven implants
were unattached in the celom. Of these, 7 were rcimplanted.
Regardless of implant source, living foreign tissue ~ v a sfound
iii the other 136 larval hosts examined.
('elomic iniplants originally pushed f ormard in the body
carity WCTC found attached either to the transverse septum
or to organs adjacent to it. Most of those mhich had heen
iiiserted into thc posterior end of the celoin and left there
hecame attached to the mesentery of thc intestine. A fev7
implants established connection with the body wall.
One type of cell found in most of the implants ohsei-ved
is characterized hy the presence within its cytoplasm of
pigment g i ~ ~ n u l e(fig.
s 3 ) . I n stained sections this pigment
appears hlack, whereas in the unstained condition, it is
y e l l o ~ ~ i i s h - h r o ~The
~ i i . granulcs vary in size froni thow too
small t o nieasixre accurately with an eye-piece inicroineler
up t o about 2.25 in diameter. These cells contain varying
aiiiounts of pigment. Some a r e so packed that they a r e
opaque. Cytoplasmic processes a r e exhibited by some of the
lightly pigmented ones.
The pigmented cells a r e not characteristically found in
specific locations v i t h respect to neighboring elements in the
implant. Ti1 some implants they are the major constituent,
566
J O H N E. ALLISOTS
whereas in others they are scattered singly or as isolated
groups throughout.
1Iistogeizesi.s of implants. All normal and overripe implants carried by Rana pipiens larvae less than 20 days
have a central core of yolk granules. One host carried a
primary iniplant of an 18-hour overripe embryo f o r 4 days
in the celoni between the body wall and a niyotome. The presence of this foreign tissue caused an inflammatory reaction
as manifested by engorgement of the vessels with blood
between the implant and the skin of the host, and by a pushing
out of the host epidermis over this area. The cells of the
host epidermis over the implant hypertrophied and also
underwent hyperplasia (fig. 4). P a r t of the ectoderm of the
implanted embryo is recognizable. The implant did not establish a vascular or tissue connection with the host. Another
embryo, stage 21, froin an overripe egg was in one Rana pipiens larval host 10 days and was resting in the celom
between the body wall and a myotome. This primary implant
is made up of a central mass of yolk surrounded by a layer
which contains mesenchyme cells and branching fibers. Pigment is present throughout as scattered granules. I n a region
adjacent to the point of contact between implant and host
(fig. 5) can be seen many pigment-filled cells of the type
described in the general observations above. Blood vessels
enter the implant here also, but do not extend beyond this
region. Peritoneum from the host forms a capsule around
the implant (fig. 5). lfitoses can be seen in the mesenchyme
cells (fig. 6). The two growths just described were the only
ones taken from larval hosts less than 20 days after implantation.
All primary normal and overripe implants that form an
attachment to, but are not imbedded in the host tissue, and
have been present within the host f o r about 20 days or longer
contain an abundance of blood vessels which are caontiiiuous
with those of the host.
A J-olk mass \\-as present in three out of 4 primary overripe
implants that had been in the host f o r 21 to 30 days (fig. 7).
G R O W T H O F A H S O R M A L FROG T R A N S P L A N T S
567
One of these implants was connected to the transverse septum,
one t o the body wall and liver, one t o the mesentery of the
intestine, and one between a myotome and the body wall.
One out of three from overripe eggs which had been established in primary hosts from 31 to 40 days contained a
yolk mass. Two of these were in the mesentery of the intestine and the other one was attached to the transverse septum.
The pigment cells are more numerous in these older implants.
Of the normal and overripe implants retained in Rana pipiens larvae longer than the ones described above, and of those
that were reimplanted, only 10 contain an aggregate of yolk
granules. I n all but two, these aggregates are relatively
small.
The implants in 5 of 13 Rana catesbeiana larvae contain a
large yolk mass. One of these was in its host 82 days, one for
96 days, two for 104 days, and one f o r 171 days. Two of
these implants had become attached to the transverse septum, two to the mesentery of the intestine, and one t o the
body wall. The yolk masses are surrounded by a layer of
typical dense connective tissue, the cells of which resemble
connective tissue cells of the mesentery of the host and thus
could have been of host origin.
The remaining normal and overripe implants into Rana
pipiens larvae are made up primarily of pigmented cells
interspersed between varying amounts of connective tissue. I n
some, this is so sparse as t o be almost indiscernible. I n others,
the fibers merge into hyaline deposits in which cells are trapped. Such deposits resemble intramembranous bone.
These results, although in some cases based on observations
made on one animal, point to the following course of events
in the histogenetic picture of implants of Rana pipiens embryos into larval hosts of the same species: There is an
inflammatory reaction in the host tissues surrounding the
implant immediately following implantation. Rossle ( '43)
and Waechter ( %I), in discussing a similar observation, contend that such a response is a defensive reaction on the
part of the hosts which tends toward destruction of the
,568
J O H N E. ALLISON
implant. l\Titschi ( '52) suggests that iiiflaininatioii of this
type may be due to a n incompatibility between host and
implant arising a s a ieesult of their difference in age. This
inflammatory reaction is accompanied by some destruction of
the organized embrj-oiiic implant tissues, and by a breaking
down of the host peritoneurn at the region of contact between
the iniplant and host tissue. These events a r e followed by a
reorganization of the implant, i n most cases into g~*owths
characterized by the presence within them of pigment-filled
cells. I n most implants the yolk is resorbed within 40 days
following implantation.
Since 5 of 13 implants of embryos from overripe eggs into
Raiia catesbeiana larvae still retain a considerable mass of
yolk a f t m having been in the host 82 to 171 days, it would seem
that a species factor is involved in this type of heteroplastic
cschaiige such that it prevents a resorption of the yolk.
Iwplants contailzing organ parts. Readily identifiable organ parts, such a s notochord, cartilage, boiie and nerve tissue,
were observed in 6 primary implants from overripe eggs,
three primary implants from riormal eggs and one secondary
implant fi.oiii a nornial egg. Examples a r e shown i n figures
iand S.
It is assumed that differentiated organ parts in the implants arise f i'om organ primordia riot eliminated during the
initial process of destruction described above. This assumption is supported by the observation that 9 out of 10 implants
producing well defined organ parts mere primary implants.
Apparently in most cases any organ primordia present in
the primary implants a r e resorbed or hcconie disorganized
-1iortly after implantation.
Of 136 implants to larval hosts examiiied, 119 ~ W I Y from
overripe eggs and 50 from normal eggs. Analyzed statistically, the ratio in lai-ral hosts of implants from overripe
einhryos t o implants from normal embryos which contain
organ parts does not differ significantly from the ratio of all
the overripe implants to all the normal implants i n larval
hosts observed. Thus overripeness has 110 apparent effect
G R O W T H O F A B S O R M A L FROG T R A S S P L A N T S
569
on the suppression or production of organ parts in the
implants.
The very small number of implants into larval hosts which
exhibit organ parts when compared with the total number
studied leads to the assumption that the implanted tissue
though remaining viable, apparently through influences exerted by its host environment, loses its ability to develop in
a n organized manner and instead ashumes the rolc of a n
uncoordinated mass of growing tissue.
Metastascs and irzfilfration. I n 54 hosts which contained
implants from overripcl embryos and in 26 which contained
implants fi-on1embryos from riorrrlal eggs, pigmented growths
of iinplant origin a r e associated with the mesentery of the
intestine. These are found scattered along the vascular system
a s nodes of variable size (fig. 9 ) . Some a r e also seen scattered
over the surface of the peritoneal covering of the body and
of organs such as lieart and liver. I n other cases, pigmented
strands a r e seen between separate nodes of these growths.
Most of these follow the course of host blood vessels.
Of all other CRSPS examined, three primary implants from
overripe eggs, one tertiary implant of a n embryo from a
normal e g g and two primary iniplaiits from normal eggs
exhibited either metastases 01’ infiltration 01- hoth. X description of selected examples of these follows.
One implant of ail embryo f r o m a normal egg was carried
through a series of three larval hosts. pigmented and nonpigmented cells h a r e penetrated the muscles of the body wall
of the third host and a r e infiltrating throughout the connective
tissue beneath the skin. Some of the pigmented cells can
be seen associated with the blood vessels of the host.
Figure 10 shows what appears to be a large metastasis in
the body wall of another larval host. This is from a primary
implant of a n embryo from tin overripe egg. The blood vessel
which could have served to carry the elements of the seconcla r y growth can he seen between the two. This particular
host also has several pigmented nodules of implant tissue
along the blood vessels of the mesentery of the intestine.
570
J O H N E. ALLISON
Typical pigmented cells of another primary implant froin
a normal egg (fig. 11)has infiltrated between the liver cords.
Several small strands and a large one extended anteriorly
from the liver. These are attached to the transverse septum.
Another primary implant fuoni an overripe egg is connected to both the liver and the transverse septum. There is
no apparent infiltration of liver tissue associated with this
growth : however, pigmented cells do penetrate between the
muscle fibers and blood vessels of the transverse septum.
Peritoneum and connective tissue of the host body wall is
invaded by another primary implant from a normal egg (fig.
12). There is a great deal of infiltration along a series of
blood vessels in the host connective tissue and metastases made
up of nodules of pigmented cells are present throughout the
connective tissue on one side of this implant.
Pigment-filled cells come to form the major constituent of
most of the implants. Many of the implants increase in
volunic, even surpassing that of the original implant. Although mitotic figures in these pigmented cells are extremely
hard to find, they did occur in about the same ratio as mitoses
in the surrounding implant and host tissue, with the exception
that there are many more in the epithelium of the host
intestine.
Tmplarzts i n f o adult hosts
The histologic picture of the implants carried by adults is
similar in many respects to that seen in the larvac. At the
end of 88 days, primary implants of embryos from overripe
eggs into the orbit of four adults could not be found. They
had been completely resorbed. I n 9 other adult hosts, the
implants were either fastened to or imbedded in the ventral
body wall. The implants in 6 of these are surrounded by what
appears to be a type of dense connective tissue. Among
cellular constituents of this tissue are fibroblasts, macrophages and lymphocytes. These are interspersed among
branching fibers. Such tissue is seen to penetrate between
the muscle fibers and normal connective tissue of the host
G R O W T H O F ABNORMAL FROG TRANSPLANTS
571
(fig. 13). The region where the epidermis has healed at the
implant site is filled with a wedge-shaped mass of epidermal
cells, some of which are smaller than the normal epidermal
constituents on either side. These animals have carried
an implant for 17 days. Of three other adults, two have carried an implant f o r 37 days, the other for 46 days. These iniplants are surrounded by connective tissue. There are no
extensions of this between the muscle fibers or into the
connective tissue of the host, as there are in the animals described above.
These results suggest that excess tissue present between
muscles of the body wall around the implant in adult hosts
17 days after iniplaiitation is provided by the host in response
to the cut through which the implant was introduced.
EpitMioma-like travwplants
Of 41 animals which had carried transplants of epitheliomalike tissue for at least 24 days, 10 exhibited abnormal growths.
None of 24 hosts which carried normal grafts f o r the same
number of days were so affected.
I n the 10 hosts which showed abnormalities, the transplants
are present as outfoldings of the transplanted ectoderm continuous with the ectoderm of the body wall of the host.
Beneath the ectodermal folds is an abnormally wide layer
of connective tissue. The muscle and peritoneum of the body
wall of the host are involved to a lesser degree (fig. 14).
The ability of these epithelioma-like growths to retain their
identity after being transplanted to normal hosts is an indication that the property of uncontrolled growth is intrinsic
within their cells.
DISCUSSION
In higher vertebrates, a tumor is usually classed as malignant if it is capable of infiltration, metastis, recurrence after
removal, and transplantation. Also it has been found that
normal tissue when implanted in a host of the same o r of
572
J O H S E. ALLISON
different species frequently undergoes iiecrosis, ~vhei*eas
iiialignarit tissue will continue to thrive.
In seeking to apply these attributes of nialignancy to amphibian material, one meets with a certain difficult>- because
mther frequently homoplastic and heteroplastic grafts of
iiormal embryonic tissue grow well. However, Belogolowy
( 'IS), Diirkeri ( 'as),Lehmann ( 'SO), 8pemann ( '42)' and
Waechter ('50) and others report also infiltratioii of host
tissues by uridiff erentiated graft cells t o which thcy w.;crihc~
it maligiiant character. Since loss o f ability t o diffei*eiitiatc
is a constant character of some of the cell.; in enihi~~-os
froiir
overripe eggs (Witschi, '52), i t was decided to study side
by side the developmental potencies of cmbryoiiic tissue
from normal and from ovcrripe cggh. Such tissue when implanted to the body cavities of older larval hosts a i d adult
hosts exhibits characteristics fulfilliiig the i~cquii~ciiientsof
malignancy in higher vertebrates. A clear explanation of
the similarities in the effects on tlic host of implants f ~ o i r i
nornial and overripe eggs in this phase of the probleni is
not iii evidence ; however, results indicate that the ciiviroiiiiieiit within the host body is such a s to influence greatly
the path of differentiation of tissues placed therein. T h e w
is a possibility that abnoriiial cells of overripe cmb~*yo>,
;tlready adversely affected, may die in the unfa\-oral>lci environrnmt afforded by the body cavity of a noixial host. If
such were the case, surviving less damaged cells -\vould
l
and no qiqnificaiit
behave like a n implant of n o ~ ~ m atissue
diff ereiices between the reaction of noi-nial arid overripe
implants should he expected.
Lehmann ('SO) suggested that the lack of oxygen and lhc
excess of carbon dioxide, which implants of normal tissue
must endure while becoming established in the host, map alter
their cells in such a way as to cause them to become malignaiit.
This might offer a n explanation for the fact that infiltrating
growths were seen t o derive from normal a s well as froill overripe implants.
(+ROWTH O F A B S O R M A L FROG TRANSPLANTS
573
Thc epithelioma cxpcriments tend to support the contention
that abnormal cells from overripe embryos possess at least
one characteristic different from the cells of normal embryos,
namely that of uncontrolled growth. Excessive growth was
exhibited by transplants from overripe embryos to normal
hosts, whereas those from normal donors to normal hosts
lost their identity a s transplant tissue. I n either casc transplantation provided a n environment similar to that which the
cells would have had if they had been left in the donor. I n
this way, environment was eliminated as a factor i n trying
to determine the cixisteiicc of differences between thc two
types of tissue.
All primary irnplaiits from normal eggs and all but 10
from overripe oiies were left in the host more than 40 days.
Some were left a s long as 219 days. It is possible that more
differences might have been noted between the behavior of
overripe aiid normal material if in the implant series the hosts
had been killed at intervals during a period immediately
following implantation. Such was not done, because it was our
intention to permit the implants to remain in the host as long
a s possi1)lc in order to facilitate expression of any malignant
potencies.
SrMMARY
1. An attempt was made to compare tho developmental
potencies of embryonic tissue derived from normal and from
overripe Rana pipiens eggs.
2. In one series of experiments, such tissues were used
to make homoplastic and heteroplastic implants into the
orbit and the celom of normal larval and adult hosts. These
implants, regardless of source, became living masses of foreign
tissue in most of the hosts.
3. Following is a n outline of the histogenesis of the
implants, a s it is discussed in the text:
( a ) Initial destruction of many implant cells including
organ primordia.
(b) Vascular connection made with host.
574
J O H N E. ALLISOX
(c) Mitoses and reorganization within the implant into
typical growths characterized by the presence within them
of specific pigment-filled cells.
( d ) Gradual resorption of yolk originally present within
the implant.
4. Organ parts and other accessory structures appear in
some of the implants.
5. The ratio in larval hosts of implants from overripe embryos t o implants from normal embryos which contain organ
parts does not differ significantly from the ratio of all the
overripe implants t o all the normal implants in larval hosts
observed.
6. Implanted tissue though remaining viable, apparently
through influences exerted by its host environment, loses its
ability to develop in an organized manner and instead, assumes the role of an uncoordinated mass of growing tissue.
7 I n many instances the implants from metastases and
infiltrating growths made up primarily of pigment-filled cells.
8. Bletastases and infiltration were not observed in implants t o adult hosts.
9. I n a second series of experiments, epithelioma-like
tissue found growing on embryos from overripe eggs was
transplanted t o the prospective flank region of normal host
embryos of the same age as the donors.
Many such transplants develop into outfoldings of ectoderm
continuous with the ectoderm of the body ~ 7 a l lof the host.
This overlies a layer of connective tissue which also appears
to be of transplant origin. Similar transplants of tissue
from normal embryos show no such activity. This was interpreted as an indication that uncontrolled growth is a property
of the cells of such epi thelioma-like transplants.
LITERATURE CITED
R A C T Z M ~ N H.
N , 1929 Uber bedentungsfremdc Selbstdifferenzierung aus Teilstucken des Amphibienkeimes. Naturwiss., I 7 : 818-827.
BELOGOLOTIT,
G. 1918 Die Einvirkung parasitaren Lebens auf das sich
entwickelnde Aniphibienei (den “Laichball”).
Roux’s Arch., 43 :
555-693.
G R O W T H O F ABXORMAL FROG TRANSPLANTS
575
BIERICH,
R. 1922 Die Resultate der intrabdominalen von Kaltbluter embryonen
in den erwachsenen artgleichen Organismus. Roux ’s Arch., 50 : 593-606.
B R I G G S , R. w., AND N. J. BERRILL 1941 Transplantation experiments with a n
ectodermal growth of frog embryos. Growth, 5: 273-284.
IKRKEN,
B. 1926 Das Verlialten embryonaler Zellen im Interplantat. ROUX’S
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KUSCI-IE,W. 1929 Interplantation umsehriebener zellbezirke aus der Blastula
und der gastrula von Amphibien. I. Versuche a n Urodelen. Roux’s
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L E H n l a N X , F. E. 1950 Entwicklungsphysiologische Analyse von Teratoman.
Rev. suisse Zool., 5 7 : 13-18.
ROSSLE, R. 1943 Weitere Versuche iiber embryonale Implantationen nebst
Bemerkungen zu H. Spemanns nachgelassener Arbeit. Wirchow ’S
Arch., 311: 230-251.
SHUNWAY,
W. 1940 Stages in the normal developnient of Rana pipiens. I.
External form. Anat. Rec., 78: 139-148.
SPEMAKN,
H. 1942 o b e r das Verhalten embryonalen Gewebes im erwaehsenen
Organismus. Roux’s Arch., 141: 693-769.
T~;AECHTER, HILDEGARD
1951 Implantation von indifferentem embryonalen Gea e g e in die Leibeschohle ervvachsener Molche. ROUX’SArch., 1 4 4 : 572617.
WITSCHI,EMIL 1930 Experimentally reproduced neoplasms in the frog. Proe.
Soc. Exper. Eiol. and Med., 2 7 : 475-477.
1949 The larval ear of the f r o g and its transformation during
metamorphosis. Z. Naturf., 4 : 230-242.
1951 Hormonale Modifikation der Geschlechtsdiff erenzierung bei
Alytes obstetricans. Z. Naturf., 6b, No. 2: 76-85.
1952 Overripeness of the egg a s a cause of twinning slid teratogenesis. Cancer Rescareh, 1 2 : 763-786.
PLATE 1
EXPL.%NATION OF‘ FIGURES
1
Tlie loner two a l e embryos btage 1 7 from overripe R. pipiens eggs. These bear
epithelionia-like growths of type u s 4 in tr:tnsplantation series. The other 13
iiornial eiiibryo same stage. X 12.
2
Cross section of one abnormal animal i n figure 1.
3
C’ells filled with varying amounts of pigment drawn from a primary overripe
implant to R. pipiens larva. x 1600.
x
100.
4 Section of implant f r o m X. pipiens stage 26. Primary implant a t A from
5-day overripe egg is betiwen body wall aiitl a myotome. Enibr\o ivah
2.7 nini (about stage 18) when introduced to host stage 2 6 , and w a s
enrried three days. Note eiigorgement of blood vessels urider pel itoneuiii
of body wall, pubhirig out of ectoderm, :uid liypri trophy am1 hyperplasia
of cells of ectodc>im over implant. X 150.
3
Primary implant in X. pipiens stage 26 from 5-day overripe egg. It is
attached to the peritoneum adjacent t o a myotome. Embryo was stage 21
when introduced to host stage 26, and was carried 70 days. Blood vessels
are entering a t poiiit of attacliment. Pigment-filled cells WI e nlso present
in this region. Pigment tlirougliout r(’st of implant is piewut a i scatteittl
grnnules. x 100.
Mitotic figure a t A and surrounding are;\ are from region outlined by solid
line in figure 5. X 1500.
Primary implant in R. pipiens stagr 27
attached to transverse septum. Embryo
host htage 26 and was carried 29 clays.
and t n o masses of yolk a t I3 and C. X
8
is froiu 5-day overripe egg. It is
was stage 21 when introduced t o
Present are some cartilage at A
100.
Primary implant from normal egg in R. pipiens stage 37. Embryo was
stage 18 when introduced t o host stage 26. Implant was carried 7 7 days. Cell
inass a t A resembles brain or spinal cord. Lens a t B is surrounded by retinal
and pigmented layers of the eye. X 100.
GROWTH OF ABNORMAL FROG TRANSPLANTS
J O H N E. ALLISON
577
PLATE I
PLATE
2
EXPLANATION O F FIGURES
9 Primary implant in R. pipiens stage 30. Metastases are in mesentery of
intestine. Embryo from 4-day overripe egg was late gastrula when introduced
t o host stage 26 and was carried 193 days. X 85.
1 0 Primary implant in R. pipiens stage 31 is from 5-day overripe egg. It is
seen a t upper left imbedded in body wall. Embryo was 2.5mm long (about
stage 21) when introduced to host stage 26. It was carried 76 days.
Blood vessels at A appears t o be possible route for metastasis a t B.
Cells of skin have hypertrophied and hyperplased over the metastasis. X 100.
11 Primary implant in R. pipiens stage 30 from normal egg. Embryo was
stage 18 when introduced to host stage 26 and was carried 140 days. Lightly
and heavily pigmented cells have infiltrated the liver in the region outlined.
X 85.
1 2 Primary implant in R. pipiens larva stage 27 is from a normal egg. I t
is attached to the dorsal body wall. Embryo was stage 18 when introduced
to host stage 26, and was carried 101 days. Main part of implant a t A is
attached by a thin strand of tissue. From this point heavily pigmented cells
infiltrate along and in blood vessels toward skin. Skin is greatly fnlded
over this growth. X 150.
13 Secondary implant in body wall of R. pipiens adult is from 5-day overripe
egg. Embryo was 2.8mm long (about stage 21) when introduced into first
host. This host was stage 26 when embryo was introduced and stage 30
when killed 99 days later. Implant was attached to transverse septum. Tinplant had been carried 17 days by the adult. Cone shaped region a t A is
possible place where cut in epidermis has healed. There is a dense connective
tissue band a t B which penetrates through muscle t o the celom. This possibly
is tissue mobilized by host in response to wound. X 100.
14 Section of body wall through region of epithelioma-like transplant from
R. pipiens stage 24. Note excessive folding of body wall. X 100.
578
GROWTH OF ABNORMAL FRO0 TRANSPLANTS
J O H B 1. A L l d H O N
579
PLATE 2
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