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A VERY YOUNG OVUM
IN SITU
PROF.Cfc.l LEOPOLD
GEHEIMER MEDIZINALRAT, DIRECTOR OF THE ROYAL GYNECOLOGIC
CLINIC AND SCHOOL FOR MIDWIFERY. MEMBER OF THE
ROYAL MEDICAL BOARD OF DRESDEN.
COMPRISING THE FOURTH VOLUME OF THE "ARBEITEN
AUS DER FRAUEN KLINIK IN DRESDEN."
WITH SIXTEEN LITHOGRAPHIC PLATES
AUTHORIZED ENGLISH TRANSLATION
BY
W. H. VOGT, M. D.
GYNECOLOGIST AND OBSTETRICIAN TO THE LUTHERAN HOSPITAL
ST. Louis, MISSOURI
C. V. MOSBY CO.
ST. LOUIS
1907
Privilege of Copyright reserved by
S. HIRZEL, Leipzig, Germany,
Oct. 10th, 1906. Act of March 3rd, 1905.
CONTENTS
PAGE
Introduction - - 5-g
Consideration of the Reconstruction of the Ovum From
Section 1 to 160 - 10-33
CHAPTER
I. The Mucosa Elevation, which Harbors the Ovum,
and the Decidua Vera - - 34-42
II. The Embedding of the Ovum. The Capsularis.
The Fibrinous Cover ("Gewebspilz" of Peters) 42-51
III. The "Eianlage" and the Ovular Chamber.
("Eikammer") 51-54
IV. The Trophoblast and its Surrounding Blood Spaces.
The Intervillous Circulation 54-64
V. The Syncytium 64-66
Explanation of Figures 67-69
101276
INTRODUCTION.
The Ovum which forms the subject of this investigation orig-
inates from a young woman who committed suicide with phos-
phorus. The post mortem examination was performed by -Ober-
medizinalrat Prof. Dr. Schmorl, in the pathological institute . of
the Friedrichsstaedter hospital in Dresden.
The fact that this woman had committed suicide suggested
the possibility of a pregnancy and for that reason the anterior
wall of the uterus was split and the specimen carefully hard-
ened, first in formalin and then in increasing strengths of alco-
hol. Although repeated and careful examinations of the endo-
metrium were made with a magnifying glass, no ovum could
be detected. Still the greatly swollen mucous membrane which
was divided into irregular areas by furrows, as we are accus-
tomed to see in a pregnant uterus, was conspicuous and induced
us to make repeated examinations during the process of harden-
ing. Finally a small spot was detected, lighter than the sur-
rounding tissue, situated on the posterior wall of the corpus uteri
a little below its center and immediately above a deep furrow.
In order to study this small spot more carefully, a square piece
of mucous membrane about 1 cm. long, with its underlying mus-
cularis was excised and after further hardening by the labora-
tory assistant, Mr. Ernst Thomas, cut into a continuous series of
sections, five microns thick and then stained in haematoxylin-
eosin. The middle one hundred and sixty sections comprised a
very small ovum whose measurements were 1.4 mm. long 0.9
mm. deep and 0.8 mm. wide, an ovum about the size of that
described by Peters, however a trifle smaller, its length being
0.2 mm. shorter than that of Peters, which measured 1.6 mm. in
length, 0.8 mm. in depth and 0.9 mm. in breadth. No information
could be obtained concerning the time of the last or first missing
menstruation. Hence it seems useless to attempt any estimate of
the age of this ovum. In the following pages it is intended to
compare the results of the microscopic examinations with those
of Peters, since this one and the one described by him are the
two smallest and youngest human ova ever observed.
5
D INTRODUCTION.
The accompanying plates were carefully drawn by the artist,
Richard Scholz, exactly true to nature and without any attempt
to obtain a diagramatic effect.
I wish to thank the artist, as well as the publisher, for the ex-
cellent reproduction of these figures and for the valuable ser-
vices they have thereby rendered science.
There will be considered:
1st. The Decidua Vera.
2nd. The Implantation of the Ovum with the formation of
the capsularis and the fibrinous covering, which lies over the
ovum on the mucous membrane. ("Gewebspilz" of Peters.)
3rd. The "Eianlage" and the Ovular Chamber "Eikammer."
4th. The Trophoblast and its surrounding blood spaces.
5th. The Syncytium.
A germinal plate which was found in Peters' specimen could
not be discovered in any of the sections of this new ovum.
Whether its absence be possibly due to the general phosphorus
poisoning must remain an open question until more specimens
of such an early stage are obtained. It must be remembered
that the woman from whom the Peters ovum was obtained, also
committed suicide, but with caustic potash, which is capable of
producing a very acute hyperemia of the mucous membrane of
the pelvic organs.
Some of the ova described by Marchand, Rossi Doria, Graf
von Spec, Heukelom, Mertens, etc., can hardly be pronounced
normal specimens free of objections. The indications for the
post mortem in such instances will necessarily be either a suicide
or an acute illness caused by poisoning or some intercurrent dis-
ease (such as pneumonia, burns, etc.), or a disease of the sexual
organs necessitating a removal of the uterus. All of these con-
ditions, especially in the early stages of pregnancy, will hardly
ever fail to lead to some changes in the circulatory apparatus of
the uterus. Nevertheless some portions of both the uterus or the
ovum may be found in a very satisfactory state of preservation.
Of the three young ova described by Marchand, the first had
a defective fibrinous capsule, supposedly injured during a curet-
age. The second was infiltrated with blood and yet Marchand
did not hesitate to investigate these defective specimens, to study
such difficult problems as the trophoblast and the syncytium and
finally drew, although with great precaution, certain conclu-
sions.
In order to satisfy myself of the value of this specimen I sub-
mitted several sections to my esteemed colleagues, Obermedizin-
alrat Prof. Dr. Schmorl and Prof. Graf von Spec in Kiel, whom
INTRODUCTION. t
I wish to thank for a careful examination of these slides. Prof.
Graf von Spec did not question in the least the good histological
preservation of this specimen. He called attention to the great
scientific value of this new ovum and pointed out the frequent oc-
currence of a congested area surrounding the seat of implantation
in this stage of development. The ovum of Peters, which may
be one-half to one day older, also shows this marked congestion
of the blood vessels in the immediate neighborhood of the ovum.
We must bear in mind that during or immediately after the
penetration of the ovum into the endometrium, a pronounced af-
flux of blood is a natural consequence. Regarding this only a
study of other very young ova, which can be expected in the
near future, will throw more light on this problem, until then
it seems permissible to disregard the objection that the poison-
ing with phosphorus might interfere with the reliability of the
histological picture presented by our specimen. At any rate it
must be emphasized, that this latest ovule, exactly like that of
Peters', the one demonstrated by Graf, von Spec in Kiel, and
the very small ovum described by me in my atlas ("Uterus und
Kind") were found in' situ embedded in an uninjured mucous
membrane in a perfect condition.
The detailed discussion in the following chapter will be simpli-
fied by a general preliminary consideration of the mucous mem-
brane elevation which harbors the ovum. This is done in the
following table, which contains only the findings pertaining to
this point in the sections 1 to 160.
This table shows plainly that the fibrin coat ("Gewebspilz" of
Peters) which covers the top of the mucous membrane elevation
("Schleimhauthuegel") consists of two adjoining portions. The
one we can follow from sections 10 to 68, where it has almost dis-
appeared. The second portion begins in section 75 and disap-
pears again completely in section 103. In a similar manner the
glands and the blood vessels can be followed. This preliminary
general consideration furnishes a good picture of the enormous
number of glands and capillaries and shows how the glands
gradually are crowded out through the dilatation and rupture
of the capillaries in the surrounding tissue.
This general survey alone demonstrates the fact that the mi-
nute ovum is completely encircled by free blood spaces.
It affords me great pleasure to say a few words regarding the
translation of my book, which Dr. Vogt has so kindly under-
taken. A few years ago Dr. Vogt was a pupil of mine who for
many months assisted me both in theoretical and practical work.
I shall owe to this translation a widespread acquaintance with
8 INTRODUCTION.
this highly interesting specimen, herein described, among my
many friends and former pupils, as well as my colleagues on the
other side of the ocean.
It is hoped that this book will instigate further research con-
cerning the earliest stages of human existence.
PROF. DR. LEOPOLD.
Dresden, Koenigliche Frauenklinik, January, '07.
TRANSLATOR'S PREFACE.
The deplorable fact that numerous important contributions to
medical literature," appearing in the German language, remain
unread by the English-speaking fraternity as the result of an im-
perfect acquaintance with that tongue, induced me to undertake
the translation of this valuable study of one of the youngest
human ova found in situ. When requested to do this work by
my esteemed former chief, Prof. Dr. Leopold, I was impressed
with the necessity of preventing the English-speaking profession
from remaining unacquainted with a work of such high order.
This book should prove interesting and instructive not only
to the embryologist and the obstetrician, but also to the profession
in general, and if by its translation I shall have succeeded in dis-
seminating a clearer knowledge of this complex subject, I shall
feel amply repaid for my labor.
In conclusion I wish to thank my friend and colleague, . Dr.
Hugo Ehrenfest, for the very kind and valuable assistance which
he rendered me in the translation of the book, and I also wish
to express my indebtedness to Mr. Geo. Heithaus, stud. med..
for his aid in the reading of the proofs.
W. H. VOGT, M. D.
St. Louis, Mo., September, 1907. 4977 Lotus Ave.
10
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTIONS 1-160.
No. of
Section.
Mucous i
Without
fibrin cover.
nembrane elevation narbonn
With
fibrin cover.
y the ovum.
Epithelium
above.
Epithelium
beneath?
I.
3-
.
4-
like i and 3.
6.
The mucous
membrane
protuberance
which later be
elevation, enci
um.
comes the ovular
rcled by epitheli-
7-
Mucous membrane becomes
thicker.
Epithelium
varying in
thickness.
8.
Epithelium
varying in
thickness.
Q.
The well pre-
served surfact
which a large g
over the elevat
; epithelium in
land opens passes
ion.
IO.
Suddenly a thin fibrin
cover appears over ele-
vation ("Gewebspilz"
Peters'.)
Under which a
trace of sur-
face epitheli-
um is to be
. seen.
II.
Firbin cover increases in
thickness toward right.
Over fibrin
cover epithe-
lium varying
in thickness.
Not clear.
12.
Present.
11
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
In the small elevation on the surface of
the mucous membrane, glands and
capillaries plentiful; glands with per-
fect epithelium.
i.
Capillaries already
considerably di-
lated.
Glands as in i.
3-
Numerous capillar-
ies in elevation of
mucous mem-
brane surface.
Greatly dilated
glands with per-
fectly preserved
epithelium.
Fig- 3-
4-
like
i and 3.
6.
Blood and capil-
laries present.
No glands in sum-
mit of ovular ele-
vation. Dilated
gland in depth of
section.
Here the first suspicion of
Eianlage by presence of
syncytial cells. Accumula-
tion of leucocytes.
7-
Dilated capillaries
in elevation and
communicating bl<
ed with syncytial c
X)d spaces infiltrat-
,ells.
Scattered syncytial cells,
accumulation of leuco-
cytes.
8.
Blood spaces in ele-
vation, whose in-
tervening walls
contain fibrin.
A large gland
opens into eleva-
tion on the same
well preserved
surface epitheli-
um.
Scattered syncytial cells.
Tissue of elevation some-
what infiltrated with blood.
Q.
In the mound con-
fluent vessel
spaces.
Infiltrated with trophoblast
processes with a layer of
syncytial covering.
Fig. 4.
10.
Capillaries become
dilated in eleva-
tion.
Glands in eleva-
tion and sur-
rounding, num-
erous and plain-
ly visible. .
The inner blood space in the
ovular elevation surround-
ed with syncytial cells.
1 1.
Blood spaces as in
1 1.
12.
12
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Mucous membrane elevation harboring the ovum.
No. of
Section.
Without
fibrin cover.
With
fibrin cover.
Epithelium
above.
Epithelium
beneath?
14.
Present.
Band of epi-
To the left and
thelium on
under the fibrin
the fibrin
covering sur-
15-
Present.
cover which
face epithelium.
sinks into the right somewhat
swollen end of the fibrin cover
like the mouth
of a gland.
Instead of as
above, there is
1 6.
Present.
found only a
slight funnel-like depression of
tissue cells anc
blood cells.
Still somewhat
'7-
Present.
of a funnel-like depression of tis-
sue cells.
Somewhat of a
"
depression cov-
1 8.
An entirely structureless
ered with epi-
mass.
thelial-like
cells.
To the left un-
der the fibrin
cover, epithe-
iQ.
' "
funnel-like dep
lium present,
ression still pres-
ent. In it or rather between it
and the fibrin cover many blood
cells.
Under the fibrin
Present.
cover a long
epithelial like
band, which
At certain parts aggrega-
probably originates from the
20.
tion of leucocytes. Oc-
syncytial offshoots which closely
casionally large syncy-
approach the fibrin cover and
tial cells, but without a
may easily be mistaken for uter-
nucleus.
ine epithelium.
13
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
Trophoblast with syncytial
offshoots.
Blood present.
14.
Decidual cells between the dilated capil-
laries and glands.
Processes of trophoblast with
offshoots.
Fig- 5-
15-
1 6.
At the periphery of the ovu-
lar chamber, syncytial cells
found.
>7-
On the border of
the various cap-
illaries syncytial
cells, partly aris-
ing from endo-
thelium.
18.
Same.
Numerous elonga-
ted and some
greatly dilated
glands.
Same.
Fig. 6.
19.
Trophoblast with syncytial
offshoots pass well up un-
der the fibrin covering.
20.
14
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
Mucous membrane elevation harboring the ovum.
Without
fibrin cover.
With
fibrin cover.
Epithelium
above.
Epithelium
beneath?
Present.
Becoming steadily longer
and lies like a worm with
its thick end on the right
of the ovular elevation.
Epithelium
varying in
thickness on
the fibrin cov-
An epithelial-
like band com-
ing from both
sides under the
fibrin cover.
Most likely syncytial cells.
23.
24 and 25.
In elongated part, connec-
tive tissue arrangement.
Leucocytes, and in the
club end fibrin only. Be-
tween these two parts as
well as towards the ovu-
lar elevation many red
blood cells.
Epithelium Under the fib-
varying in rin cover long
thickness on streaks of epi-
the fibrin cov- thelial-like
er. cells.
Close on to these the long
stretched syncytial cells with
large nuclei reach, which come
from the trophoblast.
The elongated part be-
comes more and more or-
ganized, the club end is
still fibrinous but is al-
ready infiltrated with
white blood cells.
Epithelium
varying in
thickness
above.
The same.
26.
The fibrin cover becomes Epithelium
constantly longer and ' varying in
has at its right end a but- thickness,
ton-like thickening con-
taining many red blood cells. The left end
a structureless elongation. Therefore only
the middle part is organized.
As in 23.
27.
Fibrin cover very much
elongated, on the right
end knob-shaped, folded
like a corpus luteum, in
the center of fold, blood
cells. The left end sends
out an offshoot and be-
comes steadily longer.
Epithelium
varying in
thickness
fi b r i n
An epithelial-
like band un-
der fibrin cov-
er. Under the
elongated left
o n
cover.
end of the fibrin cover this band
contains a few cells with a dis-
tinct nucleus and under the
same, running parallel with it,
are seen the large syncytial
cells.
28 and 29.
same as
27-
30.
Fibrin cover as in 27.
Blood found in fold of
knob end.
At the base of fibrin cover are found syncytial
clubs with many nuclei. Close by, an epithe-
lial-like band.
Epithelial-like
band to left
under fibrin cover. Probably
having some connection with the
giant syncytial cells.
15
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
Vessels dilate, their
walls become
thinner and rup-
ture. Blood es-
capes and passes
into the neigh-
boring glands.
Eianlage, with projection,
now seen cut laterally.
Fig- 7-
22.
Endothelium of the
capillaries en-
larged. Contains
large nuclei.
Glands to the side
of the ovum may
be seen almost in
their entire
length. Various
others filled with
blood.
Eianlage has a projection
with a broad end. Ovular
chamber filled with blood.
23.
"
"
Eianlage with trophoblast
buds and projections which
anchor themselves in the
periphery of the ovular
chamber. Projections with
trophoblast nuclei and syn-
cytial covering.
24 and 25.
At the broad base
the blood vessels
become more con-
fluent and take up
the protruding
trophoblast proc-
esses.
To the left and
right of ovum ex-
cellently pre-
served, some bro-
ken open and
filled with blood.
Eianlage, larger and takes on
a more definite form.
26.
Toward the ovular
chamber the con-
fluent blood ves-
sels are now divid-
ed by an arch-like
arrangement of
fibrin.
Eianlage as in 26, passing
from it to the fibrin cover
are five trophoblast proces-
ses with offshoots of syncy-
tium. In the surrounding
of the Eianlage decidual
cells.
Fig. 8.
27.
Passing from the Ei
blast processes and s
free blood, likewise i
anlage to the knob e
yncytial offshoots are
n the fold of the fibr
Eianlage manifold, elongat-
ed and somewhat com-
pressed by free blood,
nd of the fibrin cover, tropho-
seen. Between the offshoots
n cover.
Section
26-28.
Fig. Q and
Fig. 10.
Section
3-
Fig. n.
28 and 29
same as
27.
30.
16
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Mucous membrane elevation harboring the ovum.
Section.
Without
fibrin cover.
With
fibrin cover.
1
Epithelium
above.
Epithelium
beneath?
Between fibrin cover and
3>-
periphery of the ovular
chamber, many processes
of syncytium with many
nuclei.
32.
Between fibrin cover and
neighboring border many
syncytial giant cells.
Band of epithe-
lium not dis-
tinct.
To the right un-
der the fibrin
cover, presum-
ably a rem-
nant of epi-
thelium.
33-
Present.
Below and above fibrin cover no
band of epithelium left.
34-
Giant syncytial cells reach
to fibrin cover.
Absent.
Only toward
and under fib-
35-
Present.
rin cover are 4
epithelial-like
formations.
36.
Present.
Elongated with club-
shaped end.
Is similated by
rows of epithe-
lial-like syncy-
tial cells.
Present.
Under the knob
37-
.
Elongated with knob end,
which appears like a
thick fibrin band.
Band of vary-
ing thickness.
end an epithe-
lial-like band
surrounded by
blood cells.
The elongated part of the
fibrin cover here resem-
38.
bles some of the previous
specimens. The knob-
like end is more fibrinous
but is already beginning
to be infiltrated with
,
leucocytes.
17
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
Tropholbast processes. Coat
elevated by blood and con-
sists of two layers. Row of
nuclei with syncytial cov-
ering.
31-
Trophoblast processes with
offshoots pass toward per-
iphery, where fibrin cover
lies. Coat elevated, 2 lay-
ers: Langhans' cells and
syncytial covering.
Fig. 12.
32.
Eianlage with partly elevat-
ed coat.
33-
34-
Blood vessels dis-
integrating.
Processes of trophoblast.
35-
Mesoblast and Ectoblast
(Trophoblast and Syncytial
cells) pass under entire fib-
rin cover along the wall of
the ovular envelope.
36.
Owing to the in-
creasing size of the
ovum the sur
rounding blood
vessels become
more and more
disintegrated,
to the ovum but doe
ovular chamber.
To the right and
left of ovum two
glands. In the
middle of these
a third gland
which gradually
dilates and passes
s not open into the
Fig. 14.
37-
The same.
The syncytial cells pass up
to the ovum periphery and
make the impression of sur-
face epithelium.
38.
18
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
Mu
Without
fibrin cover.
:ous membrane elevation har
With
fibrin cover.
boring the ovum.
Epithelium
above.
Epithelium
beneath?
39-
the one side of wh
arising from the t
Present.
Under the long offshoots
of the fibrin cover a
long blood vessel present,
ich consists of broad cells
'ophoblast.
Over the knob
end a worm-
like cover.
40.
i
The elongated part of the
fibrin cover is made up
of distinct tissue. The
knob end is arranged ex-
actly like a corpus lute-
um.
Over the knob
end a band
varying in
thickness.
Under the knob
end reaching
from the bor-
der, an epithe-
lial band.
41 like
40.
Same.
Under the entire fibrin
cover, large svncy tial cells
are seen.
42.
Under the knob end syn-
cytial giant cells.
43-
The knob end of the fibrin
cover is infiltrated with
leucocytes.
45-
Under the fibrin cover the
row of syncytial cells
have become consider-
ably larger.
47-
'
Present.
49-
Present.
gated syncyti
might be mist
epithelium.
Under the fib-
rin cover elon-
al cells, which
aken for surface
5-
The knob end becoming
shorter and rounder.
Elongated syn-
cytial cells be-
low.
19
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160
Blood
vessel.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
Left gland with
broad opening.
Right gland surro
die one which reac
with blood.
unds entire ovum. The mid-
hes to the ovum becomes filled
Fig. 13.
39-
From the trophoblast an ex-
tension of syncytial cells
pass to the epithelial-like
band on the border under
the knob end.
40.
As in 40.
41 like
40.
Covering with syncytial buds.
42.
43-
of the ovum can be
the entire length tc
The two glands to
the right and left
followed almost in
their openings.
Covering has two rows.
45-
A broad part of the
contact a double ce
visible. Here as in
which anchor them;
ovum lies under the
1 row (Langhans* eel
48 one' sees large of
elves into the peripri
Processes of trophoblast with
offshoots,
fibrin cover. At the point of
,s and Syncytium) are plainly
"shoots, from the trophoblast,
ery of the ovum.
Fig. 1 6.
47-
Offshoots with covering
which become anchored.
(Section 50.)
49-
In the periphery of
the ovum toward
the middle of the
mucosa uteri, an
enormously dilat-
ed vessel.
Eianlage with offshoots as
heretofore.
5'-
20
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160
No. of
Section.
Mu
Without
fibrin cover.
cous membrane elevation hai
With
fibrin cover.
boring the ovum.
Epithelium
above.
Epithelium
beneath?
52.
Under fibrin cover enor-
mous syncytial cells.
53-
The fibrin cover becomes
more compressed. (Al-
most structureless.)
Giant syncytial cells be-
low.
54-
The knob end rounder and
thicker, under the same
thick syncytial cells, with
many nuclei.
55 and 56.
Under the fibrin cover the
syncytial cells constant-
ly increasing in number.
57-
Under entire length of fib-
rin cover larger and
smaller syncytial cells are
found.
Over knob end
of fibrin cov-
er a band of
epithelium
varying in
thickness.
58 and 59.
Fibrin cover compressed,
shortened and thickened.
Beneath syncytium.
60.
structureless knob
has become organi
syncytial cells are
The fibrin cover now pre-
sents itself as an almost
the rest of the fibrin cover
zed. Under this cover the
constantly multiplying.
6 1 and 62.
Among these syncytial
cells trophoblast cells are
observed.
63.
The knob end like a but-
ton. Under the same
rows of syncytium, which
pass almost to the outer
border of the covering.
f 21
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Einlage.
Illus-
tration.
No. of
Section.
Band of ectoblast with Lang-
hans' cells and syncytium.
In Eianlage peculiarly ar-
ranged cells.
Embryonic rudiment?
Fig. 26.
52.
Eianlage lies plainly on the
periphery with a covering
of two layers.
53-
The enormously di-
lated blood ves-
sels as in 51.
Trophoblast processes with
syncytium anchored to per-
iphery of ovum.
54-
55 and 56.
Dilated capillaries.
well preserved unc
due to the confluen
Enormously dilated
present in 51.
Beautifully ar-
ranged, partly
ergoing dissolution
t blood vessels,
blood vessels still
The albuminous mass in the
Eianlage compressed by
blood in the ovular cham-
ber.
57-
To the left and
right of ovum,
glands with open-
ings.
Eianlage with trophoblast
processes and syncytial off-
shoots.
58 and 59.
The enormously di-
lated blood ves-
sels divided into
three spaces by
the decidual tis-
sue.
Eianlage with trophoblast
processes. Syncytial cov-
ering.
Fig. 17.
Fig. 27.
60.
The two glands to
the left and right
are distinctly
visible.
Eianlage, as in 60.
6 1 and 62.
The same.
Eianlage with trophoblast
and offshoots.
Fig. 1 8.
63.
22
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
Mu
Without
fibrin cover.
cous membrane elevation hai
With
fibrin cover.
boring the ovum.
Epithelium
above.
Epithelium
beneath?
65 and 66.
The knob-like end becomes
flatter.
67 and 68.
The knob becomes still
flatter and is hardly vis-
ible in 68.. Beneath it a
layer of syncytium.
69.
The knob but a small
structureless fibrinous
elevation. Layer of syn-
cytium beneath it.
70-73.
The knob is as 69. Be-
neath it syncytium,
which might be mistaken
for uterine surface
epithelium.
In 73 a layer of
epithelium
varying in
thickness.
*
74-
The knob as in 73.
As in 73.
75-
becomes longer an
cover.
The fibrinous structureless
mass, which has been
but a tiny knob, again
d lies on the organized fibrin
76-79.
The fibrinous structureless
mass becomes steadily
longer. Syncytium is
seen beneath this mass.
A band of epi-
thelium vary-
ing in thick-
ness.
Similates uter-
ine surface epi-
thelium.
80.
The structureless mass be-
comes steadily longer and
begins to appear as be-
fore.
(See sections 10-25.)
An epithelial-
like band.
Si.
82 and 83.
84.
Fibrin cover with a band lying above it.
A few cells be-
neath. (Epi-
thelial cells?)
23
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160-
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
65 and 66.
Glands plainly vis-
ible to the right
and left.
Eianlage toward the sum-
mit with trophoblast.
67 and 68.
Glands plainly vis-
ible to the left;
to the right they
are displaced.
Eianlage with beautiful tro-
pholbast processes and off-
shoots with syncytium.
Fig. IQ.
69.
The same.
The same.
70-73.
Numerous syncytial cells
scattered in the surround-
ing of the ovum.
74-
The same.
Eianlage with trophoblast
processes.
Fig. 20.
75-
'
76-79.
The same.
The same.
Fig. 21.
80.
Polymorphous and pressed
together by Eianlage and
ectoblast. The latter ar-
ranged like numerous
prongs. Trophoblast proc-
esses.
(View
of
entire
ovum.)
81.
Large trophoblast processes
with syncytial offshoots
extending to the border
and walls of the blood ves-
sels.
82 and 83.
Trophoblast processes with
syncytium plainly visible,
In the periphery of the ovu-
lar chamber large syncy-
tial cells.
84.
24
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Section.
Without
fibrin cover.
With
fibrin cover.
Epithelium
above.
Epithelium,
beneath?
85.
Under fibrin
cover a chain
86
and
87.
The fibrin cover becomes
longer and on its right
end has a knob - like
thickening.
Epithelium
spreading
above it.
.of cells (Epi-
thelium?) At
the right bor-
der a distorted
gland. Blood
has entered
it.
88.
band a fine layer ol
ed syncytial cells c
Fibrin cover larger.
Toward the epithelial
fibrin is seen;- then elongat-
ccasionally trophoblast.
A long band
of epithelium
over it.
Under it a torn
epithelial band
however in
better pres-
ervation.
Under the fib-
rin cover, os-
89.
Present.
tensibly an ep-
ithelial layer.
Present?
90-92.
Elongated.
Present.
Immediately
below it are
biids of syncy-
tium.
93-
Larger.
4
Epithelium above and beneath
the fibrin cover.
94.
Distinct.
95 and 96.
The same.
Present.
Present.
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.'
No. of
Section.
To the right be-
tween the ovular
chamber and the
adjoining glands
new capillaries
are constantly be-
ing opened.
The large gland to
the left of the
ovum reaches to
the border. The
almost in its enti
becomes lost th
periphery.
inland to the right can be seen
re length. The middle gland
e nearer it approaches the
85.
Confluent capillar-
ies at the border
becoming dis-
solved.
Also from the endott
near the border ol
cells are seen prol
syncytial cells.
Long glands to
the right and left.
The middle one
does not quite
extend to the
ovum.
elium of the vessels
the ovum cavity,
ferating, similating
86 and 87.
The offshoots of a long tro-
phoblast plug radiates
toward a vessel.
88.
etrates an adjoin
wall has been pen
its neighbor. Lar
at other places.
The syncytium passes from
a trophoblast plug and pen-
ing vessel. After the vessel
etrated it communicates with
ge clusters of syncytial cells
89.
The three glands
as in 86 and 87.
border. The sync
group of vessels in
corroded by two s
Long trophoblast processes
with syncytium at the ends.
Becomes anchored at the
vtium corrodes a vessel. A
the border are likewise being
yncytial offshoots.
Fig. 28.
90-92.
Vessels in dissolu-
tion in the border.
The glands to the
right are seen in
their entire
length. Like-
wise the left. The
middle glands
disappear.
Long trophoblast processes
anchor with syncytium.
Fig. 22.
93.
The middle gland
becomes lost
toward the bor-
der of the ovum.
Mesodern distinct and poly-
morphous. At the border
large trophoblast plugs with
syncytial covering become
anchored to the ovum per-
iphery.
94-
The middle gland
at the border of
the ovum with
large epithelial
cells.
Trophoblast at the border of
the ovum (near the fibrin
cover) has two hook-like
plugs between which is
found considerable blood.
Fig. 23.
95 and 96.
26
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Mucous membrane elevation harboring the ovum.
Section.
Without
fibrin cover.
With
fibrin cover.
Epithelium
above.
Epithelium
beneath?
97 and 98.
Fibrin cover has sunk into
the underlying tissue.
Bands of epi-
thelium vary-
ing in thick-
Bands of epi-
thelium.
ness.
99.
Fibrin cover elongated.
Likewise.
Bands of epi-
thelium vary-
ing in thick-
ness.
100.
Present.
Fibrin strip has
a layer of epi-
thelium.
Likewise
below.
IOI.
Fibrin cover depressed.
To the left epithelium.
Epithelium (?).
beneath the
fibrin strip.
Under the fib-
1 02.
~
Small fibrinous elevation.
rinous eleva-
tion, strips of
epithelium (?)
103.
Fibrin cover en-
tirely absent.
Only a tiny particle there-
of present.
Short bands,
(Surface epi-
thelium?)
104
and
105.
Absence of fibrin
cover.
1 06
and
107.
The same.
Surface epithe-
lium fairly
well pre-
served.
1 08.
The same.
I 10
and
The same.
in.
27.
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
cells close to the ov
lost however the cl
the ovum. To th
the left i to 2 glanc
Middle gland has
large epithelial
um, which become
aser they approach
: right 2 glands, to
s.
Trophoblast processes with
syncytium.
97 and 98.
Three glands.
Trophoblast processes with
syncytium.
99.
Three glands.
The middle one
begins to disap-
pear.
Trophoblast processes. Syn-
cytium reaches to the bor-
der.
100.
Three glands.
Beautiful tropholbast proc-
esses.
101.
The middle gland
is seen plainly
reaching to the
ovum.
Trophoblast processes reach
almost to the border of the
ovum. Syncytial buds
reach to the under border of
fibrinous elevation.
1 02.
The middle glands
not so plainly
visible.
The trophoblast and syn-
cytium reach to the band
beneath the beginning of
the fibrin cover.
103.
The middle gland
reaches almost to
the ovum.
Fig. 24.
104
and
105.
Two new glands
are seen on the
border of a dilat-
ed vessel.
Trophoblast processes with
syncytium and giant cells.
106
and
107.
seen. In later sec
comes more plainly
length.
The end of a large
gland is plainly
tions this gland be-
visible in its entire
Trophoblast processes.
1 08.
Capillary spaces.
The gland very
distinct, the
opening however
has become less
distinct. Other
glands with their
epithelium have
become destroyec
through hemor-
rhage.
Buds of syncytium.
I 10
and
in.
28
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
M
Without
fibrin cover.
ucous membrane elevation ha
With
fibrin cover.
rboring the ovum
Epithelium
above.
Epithelium
beneath?
I 12.
Not a suspicion ol
a fibrin cover.
113.
The same.
i 14-1 i 6.
1 17 and 1 1 8.
The same.
IIQ.
A trace of fibrin present.
121
and
122.
Not a trace of fib-
rin except on
ovular elevation.
123.
and
124.
The same.
125-128.
129-131.
132.
133.
29
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessel.
Glands.
Eianlage.
Illus-
No. of
tration.
Section.
| The mouth of the
gland disappears.
In a gland filled with blood, the epithe-
lium cells are swollen, partly enlarged
and elevated from base.
Buds of syncytium.
A gland at the bor-
der. Close to the
pole of the ovum, a gland lined with epi-
thelium and greatly dilated, is seen
filled with blood.
113.
Dilated capillaries
surround the ov-
um.
A large long gland
close to the ovum
Syncytium anchor.
i 14-1 16
117
and 1 1 8.
The mouths of
two glands on
top of the ovum.
Delicate processes 'become
anchored to the periphery.
119.
Cavernous Wood
spaces in the per-
iphery of the ov-
um.
A gland opening.
Between the cavernous blood
spaces the syncytium is ar-
ranged in arches.
121
and
122.
Cavernous blood
spaces surround-
ing the ovum.
Many capillaries
open in the sur-
rounding of the
central blood
space.
Two gland open-
ings close to top
of the ovum
mound.
123
and
124.
To the right of the
ovum a gland
with a distinct
opening
alone;
capillary.
passes
a dilated
The interior of the ovum
filled with blood, this is
enclosed by arch-like ar-
ranged syncytial cells.
125-128.
A number of capil- Glands as in 132.
laries open at a
place which was
previously occupied by the nucleus of
the Eianlage and break into discern-
ible glands.
Syncytial cells become less.
In the periphery Still more syncytial cells
of the ovular ele- with large well stained nu-
vation four clei.
glands are plainly visible partly a.) dissolved in capillaries, at their
opening or in the center, b.) fairly well traceable in their entire length,
c.) not plainly visible, d.) opening not distinct, toward the interior
dilation plainly visible.
132.
Capillaries break
into glands.
Syncytial cells with large
nuclei.
'33-
30
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
Mu
Without
fibrin cover.
cous membrane elevation hai
With
fibrin cover.
boring the ovum.
Epithelium
above.
Epithelium
beneath?
134-136.
137-I39-
140.
141.
142
and
143.
The mucous membrane elevation upon which
no fibrin cover is present, shows a few conflu-
ent blood spaces in the center, besides this
beautiful glands and numerous blood ves-
sels.
144
and
145.
147.
Rests of sur-
face epithe-
lium.
151.
Divided blood spaces in the ovular elevation.
152.
' 5 1
and
55-
Surface epithe-
lium present
over ovular
elevation.
, 31
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
Syncytial cells be-
tween the blood
spaces.
Syncytial cells not so num-
erous.
134-136.
The vessels pene-
trate the glands.
i 37-' 39-
The endothelium of
the vessels contain
large nuclei, parall
lie directly on the v
;1 with these a few
essels.
140.
Numerous syncytial
cells in the sur-
rounding of the-
capillaries.
141.
Greatly dilated.
Four large glands
pass through the
ovular elevation
but much more
distinctly visible tl
b.) elongated; c. 2
In the center of the mound
many syncytial cells and
accumulation of leucocytes.
lan in 132. a.) greatly coiled;
nd d.) with distinct openings.
Fig. 25.
142
and
143.
Confluent blood
spaces in the cen-
ter of ovular ele-
vation.
Gland spaces and
mouths.
A few syncytial cells and ac-
cumulation of leucocytes in
the surrounding of the cen-
tral blood spaces.'
144
and
145.
Dilated confluent
blood capillaries in
the ovular eleva-
tion.
Glands dilated.
Between them
accumulation of
leucocytes.
147.
Capillaries greatly
dilated.
In the surrounding ol
al cells are present.
Likewise the
glands.
both many decidu-
151.
A few very large
syncytial cells
with highly stained
in the more centr
spaces.
nuclei are still found
ally located blood
152.
Central blood
spaces and with
these communicat-
ing capillaries.
I5 1
and
155-
32
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
No. of
Section.
Mu
Without
fibrin cover.
cous membrane elevation har
With
fibrin cover.
soring the ovum.
Epithelium
above.
Epithelium
beneath?
156.
Fibrin with large
ovular elevation.
nuclei passing through the
,58.
In the ovular elevaiton only the borders of the
more centrally located blood spaces can be
seen.
1 60.
Ovular elevation supplied with enormous capil-
laries.
Uterine sur-
face epitheli-
um present.
33
CONSIDERATION OF THE RECONSTRUCTION OF THE OVUM FROM SECTION 1-160.
Blood
vessels.
Glands.
Eianlage.
Illus-
tration.
No. of
Section.
To the left and
157: Centrally
large blood spaces.
right glandular
spaces partly
filled with blood.
156.
,58.
Detached capil-
laries.
Numerous glands.
1 60.
GENERAL CONSIDERATION OF THE RECONSTRUCTION
OF THE OVUM FROM SECTIONS 1 TO 160.
It may here be stated that all the microscopic sections have been
embedded with the summit of the elevation produced by the ovum,
pointing to the right, as in Fig. 2 ; thus the entrance into the fur-
row formed by the mucous membrane, also opens to the right.
In this manner the meaning of the constantly recurring terms
"right" and "left" in the description of the sections, is obvious.
I.
THE MUCOUS MEMBRANE ELEVATION WHICH HAR-
BORS THE OVUM, AND THE DECIDUA VERA.
The mucosa, which lines the corpus uteri, is about 5 mm. thick
in the fundus and anterior wall but about 8 mm. thick in the
middle of the posterior wall. As previously mentioned, numerous
deep and irregular furrows divide the mucous membrane into a
number of fields. Toward the internal os (Plate I Fig. 1. o. i.)
the mucosa suddenly becomes thinner and assumes the charac-
teristic appearance of the plicae palmatae. It is here only 2 to
4 mm. thick. Since, as is well known, the mucous membrane of
the uterus has no submucosa, a sharp line divides it from the un-
derlying muscularis (Fig. 2, g.). Any thin section passing
through both mucosa and muscularis, if held to the light, will
distinctly show this sharp division both in the pregnant and the
non-pregnant uterus. Nevertheless the connection between the
mucosa and the underlying muscularis is a very firm one. The
blind ends of the uterine glands are fastened by means of a very
fine meshed connective tissue near the funnel shaped cavities
and holes formed by the muscular fibres which are visible imme-
diately below the mucosa, both in transverse and oblique sec-
tions.
In Fig. 1 the somewhat lighter spot, marked "Ovum," indi-
cates the seat of implantation of the small ovum. It lies in a
slightly protruding elevation of the mucosa ("Schleimhauthue-
gel") nearer its slope, which gradually loses itself in a broad and
rather deep furrow, (vide Fig. 2, a 2. c. Fig. 23, sections
94 to 96.)
In about the first ten sections this little hill appears as a hori-
zontal fold of the mucosa with a sharp edge (Fig. 2. a. h.)
which, as can be seen under the microscope, begins with a blunt
34
THE MUCOUS MEMBRANE ELEVATION. 35
extremity and has a gland passing through its entire length.
This gland in its middle portion is dilated and folded and lined
in its entirety with perfectly preserved cuboidal epithelium.
Gradually this hill like elevation becomes more rounded (Fig.
2. b. h.) with a sharp groove on its right side. This shape is
maintained through almost all of the sections until section 160
is reached, where it takes on a more club shaped appearance.
The furrow to its right has become somewhat shallower and now
runs perpendicular.
We might be led to believe that this elevation was brought
about by the embedding of the ovum, but the further examination
of the specimen will show that the mound is only the marginal
elevation of a mucous membrane field, lying just above a groove,
through which the ovum has bored its way.
This ovum, like the ovule of Peters, in all probability has en-
tirely sunk into the mucous membrane, so that we also find in
this case the ovular chamber completely surrounded by a de-
cidua vera which extends up into the cap which covers the top
of the ovum and protrudes into the uterine cavity. Besides this
decidual tissue the cap contains the fibrinous cover consisting
of two layers or two flat portions, while in Peters' case the clos-
ure of the small opening on the top is formed by the fibrin lid
alone, in this case the nidus of the ovum represents a com-
pletely closed cavity which on its top carries the fibrinous cover.
In investigating the histologic structure of this mucous mem-
brane elevation our attention is at once drawn to the decidual
tissue, the glands and the blood vessels, the ovular chamber and
its fibrin cover. Since the two last named structures will be con-
sidered in the next chapter, the enveloping tissue alone shall here
be described.
It is probably unnecessary to state that the younger the ovum,
the better will be the opportunity of studying the origin of cer-
tain structures such as the decidual cells, the syncytium, the inter-
villous spaces, etc.
If any doubt has still existed concerning the fact that the de-
cidual cells are nothing else but hypertrophied connective tissue
cells of the uterine muscosa and have nothing to do with the
leucocytes, this doubt is now dispelled by this specimen.
The mucous membrane enveloping the ovum is swollen and
appears edematous. Large connective tissue cells with long pro-
cesses form a delicate network. These cells contain large nuclei
which in some cases completely fill the cell body. Some cells
have two nuclei, and like Peters (1. c. page 15.), I was able to
observe signs of karyokinesis in them.
36 THE MUCOUS MEMBRANE ELEVATION.
In the vicinity of the capillaries and larger blood vessels, which
are enormously dilated and in some places ruptured, we find many
red blood corpuscles lying between the decidual cells.
In this way the somewhat exaggerated histologic picture of
hyperemia and actual suffusion of the mucous membrane (com-
pare Fig. 25. Section 142) is formed, not unlike that described
as occurring during menstruation, by the author 1 , later by.Geb-
hard 2 , and recently by Sellheim. 3
Between the blood vessels and glands the decidual tissue is
distributed partly in fine and coarse bands, partly in form of an
outstretched network and there is as yet no distinction possible
between a decidua compacta and decidua spongiosa, since we
find thick compact layers and broad bands of decidual tissue in
the superficial as well as the deeper layers of the mucosa.
Noteworthy is the wealth of glands and their course. In the
sections on either side of the ovum (Sections 1 to 30 and 120
to 160) some of the glands can be followed in their entire length
up to their entrance into the uterine cavity (Fig. 3, Section 4.
Fig 25, Section 142). At the opening and in the middle they
are often greatly dilated. Here their walls are folded, forming
larger and smaller waves which like papillae protrude into glan-
dular lumen and suggest the picture of a beginning adenoma
In the deeper portions the glands are lined with a perfectly pre-
served cylindrical epithelium which however becomes more dis-
tinctly cuboidal the closer the glands approach the base of the
peripheral wall of the ovular chamber.
Since the latter is made up of blood spaces, as will be dem-
onstrated later, the surrounding decidual tissue also is 'infiltrated
with blood, as can be seen in Fig. 25, Section 142 (Plate XV bl.).
As a result of this mucosa hemorrhage the glands in the imme-
diate vicinity of the ovum also are filled with red blood cells and
are in a state of partial dissolution exactly like during menstrua-
tion.* The epithelium loosens from the glandular walls and
disappears among the blood corpuscles. The delicate connec-
tive tissue of the gland wall becomes loosened -and is dissolved;
the further the ovular envelope moves toward the periphery, the
more the glands become displaced. Their outer walls are flat-
tened, their inner walls dissolved. Certain sections (37 to 105)
show this process very plainly. It is best observed by following
the three longest. The one in shape of an arch, reaches the
ovum and partly encircles it, the second passes to the right, the
1 Leopold, Studien ueber die L T terus Schleimhaut (Archiv. f. Gynaek Bd. XI).
2 Gebhard. Pathol. der Weibl. Geschlechtsorgane.
3. Sellheim-Nagel. Handbuch der Phyiol. des Menschen, II 1, pages 96 to 98. Figs.
38-41.
4. Nagel. 1. o. Fig. 41.
THE MUCOUS MEMBRANE ELEVATION. 37
middle one ends blindly near the ovum and as its cavity expands,
gradually disappears.
The Figs. 14 (Plate VIII) and 23 (Plate XIII) show espe-
cially the arches formed by the left and right glands. In Fig.
22 (Plate XII) we see only a trace of the end of the middle
gland, whereas in Fig. 24 (Plate XIV) the section 165, which
lies further away from the ovum proper, again shows the end
of the middle gland, which is dilated and filled with blood,
(dr.)
From this can be seen that the glands are considerably dilated
and hypertrophied. The "Eianlage" acts as a foreign body and
presses the glands toward the wall. The "Eianlage" furthermore,
together with the free extravasated red blood, destroys and dis-
solves glands and thus helps to form the intercommunicating
blood spaces of which we will presently speak. None of the sec-
tions furnished any proof for the assumption that the glands open
directly into the ovular chamber, or that the ovum entered the
mucosa through the mouth of a dilated gland. No signs were
found of any well preserved glandular epithelium or even rests
of such lining this cavity. This is furthermore made improbable
since, as we have seen before, the glands in their entirety, includ-
ing the walls and epithelium in the immediate vicinity of the .
ovum, are dissolved as the result of its growth.
The mucous membrane elevation and all the adjoining tissue
down to the muscularis, is very vascular. The structure and
course of the blood vessels during the first weeks of pregnancy
have been carefully described in a number of excellent works
and we refer to them for further information. Here only those
facts shall be emphasized which seem most important in connec-
tion with the very earliest stage of pregnancy. Since they are
dealt with only in the work of Peters, a confirmation or further
elaboration of his contribution would seem very desirable.
In the deeper portions of the mucosa, especially in the thicker
layer of the decidual trabeculae, one still meets with little arteries
twisted like cork screws. As a rule they are found cut either
transversely or lengthwise, the latter often accompanying the
glands for a long distance. On cross section the capillaries show
around their endothelial lining a second and sometimes a third
delicate ring of endothelium. The closer the ovum is approached,
the greater becomes the number of capillaries and the greater is
their distention and the extravasation of red blood cells into the
surrounding tissue. .If the delicate wavy walls of these distended
capillaries are carefully examined, it will be seen that at certain
places there are defective areas or even tears, through which the
38 THE MUCOUS MEMBRANE ELEVATION.
erythrocytes escape into the adjoining tissue. They loosen
the glands and detach the swoolen epithelium which
now lies crumbled in the gland duct. In this way a coalescence
of glands and capillaries is effected which in the neighborhood
of the ovum results in the continuous formation of new inter-
communicating blood spaces. If Figs. 7 and 8 ('Plate IV)
marked "c," are carefully studied, this process can be better
understood than describing the same. Figure 14 (Plate VIII)
is still better, in which c and c indicate the adjoining cavities
filled with blood.
The confluent blood spaces are best seen in Fig. 22 (Plate
XII) where "c" shows fine septa as the remains of the tissue
which at first separated the glands from the capillaries. As the
"Eianlage" grows these septa become still thinner and are finally
completely dissolved, so that at "c" the blood spaces are in direct
communication with their neighbors c l and c 2 , and the latter
again with the ovular chamber and its blood spaces. Attention
is here called to the fact (which later on will be dealt with in
detail), that the trophoblast processes have anchored themselves
to the walls of these spaces at c x and c 2 . Thus it is self-evident
that in the entire periphery of the ovum these trophablast pro-
cesses are surrounded and nourished by connecting blood spaces.
These conditions are seen more distinctly in section 94 (Fig. 23,
Plate XIII). At "c" a broad blood space runs almost parallel
with the glands to the ovular chamber. At c l another one passes
from the left inward and breaks up into several blood spaces
which communicate with the ovular chamber. It is best seen
at c 2 , where a dilated capillary, running to the left and upwards,
like through a slit, appears in the cavity.
A striking similary is obvious, if we compare these pictures
with those in my atlas ("Uterus und Kind" plate VI) illustrat-
ing injected intervillous spaces in the fifth month of pregnancy,
which communicate with vessels of the serotina.
It seems justifiable to deduce from Figs. 22 and 23, that al-
ready in the first days of pregnancy, as described by Peters and
myself, the capillaries, through their dilatation, their breaking
up into the surrounding tissue and their dissolution into enor-
mous blood spaces which communicate with the ovular chamber,
form the first rudiments of the intervillous spaces.
How do these statements compare with those of Peters, Graf
von Spec (1. c. pg. 2.) and the various researches made on ani-
mals, as compiled by Pfannenstiel 1 ? In this early stage Peters
has also observed decidual cells. He could not find a sharp di-
vision in the mucosa between a compacta and spongiosa.
1 Winckel. Handbuch der Geburtshilfe. Bd. I. 1, pg. 194.
THE MUCOUS MEMBRANE ELEVATION. 39
The mucous membrane elevation in which his ovum lies, pro-
trudes but slightly and still has a superficial epithelium covering
its margin. The glands are greatly dilated, lie around the ovum
meridionally and also have no communication with the ovular
chamber, the latter pressing them aside. In his sections also vari-
ous uterine glands (Plate I) are seen filled with blood, the sur-
rounding of the ovum containing an enormous wealth of greatly
dilated capillaries which in part are changed .into wide blood
spaces exactly as in my specimen. In his Plate I, a large blood
space is seen to the right and below, like an extravasation sur-
rounding almost half the periphery of the ovum, communicating
freely with those blood lacunae in which the trophoblastic pro-
cesses are directly immersed.
Although the similarity between Peters' and my specimen is
striking, still attention must be called to certain apparently ex-
isting differences. In Peters' specimen the trophoblast every-
where lies in direct contact with the inner wall of the envelope
of the ovum and an embryonic rudiment is plainly visible. In
my specimen, on the other hand, the trophoblast has not as yet
everywhere reached the inner wall. It still partly floats, in the
blood which fills the ovular chamber. In spite of repeated and
careful examinations no trace of an embryonic formation could
be detected.
Whether this floating of the trophoblast in the blood of the
ovular chamber and the absence of an embryonic "Anlage" is pos-
sibly only due to a pathological extravasation of blood between
the ectoblast and mesoderm, is still a question which, must be
decided later. This difference, which in this connection is simply
registered, is striking and might warrant the question, whether
the ovum in my specimen is not possibly a few hours younger
than that of Peters, and, whether within the next one-half day the
junction between the trophoblast and the inner wall of the ovular
envelope would not have been completed. This, however, must
'remain a mere assumption until other specimens of such an early
stage have been found.
At any rate the same vascularity which is observed in our two
specimens, is also found in the uteri of animals. "The changes
in the maternal mucosa surrounding the ovum, in the various
mammalia, are differently described. A formation of new ves-
sels in the immediate neighborhood of the chorion, often to a
great extent, is regularly found. In advanced stages noticeable
vascular changes also occur in the deeper layers of the mucous
membrane 1 ."
"Very important are the results of researches made on animals
1 Pfannenstiel in Winekel 1. c. pg. 198.
40 THE MUCOUS MEMBRANE ELEVATION.
regarding the first relation between the surface of the ovum and
the vascular system of the maternal mucosa. This relation is
established very early, both in the insectivora and the rodents.
The ectoblast thickens at the placental site by the formation of
numerous layers and lacunae soon form which become filled with
maternal blood. The manner in which the blood gains access to
these spaces is not sufficiently understood, but it is certain that
the new formation of maternal blood vessels near the ectoblast
is enormous and that from them the ectoblast lacunae receive their
blood supply."
"The maternal blood from the very beginning remains in a
closed channel and in regular circulation. Frommel has demon-
stated the same conditions in the bat. In the carnivora, e. g., the
cat, these maternal vessel walls are also preserved in the pla-
centa. The significance of this fact for human placentation is
evident. In animals the "intervillous" spaces appear very early;
they are filled with blood from the very beginning and are in con-
stant and lasting communication with the maternal blood vessels.
These same conditions no doubt also prevail in the human."
From the above description this presumption is proved a cer-
tainty, even for the earlier stages of the human ovum. '
The vascularity of the mucosa in my specimen, in its relation
to menstrual conditions, is still another subject worthy of
consideration in this chaper
.Peters estimates the age of his ovum as from 3 to 4 days by
assuming a relation between it and the menstruation which for
the first time had just failed to appear. He bases his assump-
tion on the history of the case and on the histological condition
of the decidua, which, "in every respect resembles the description
given of decidua menstrualis immediately preceding menstrua-
tion." 2
From the history Peters endeavors to show that on the day of
her suicide (October 1st) the pregnant woman was about to
menstruate. From the state of development of the Eianlage and
the entire ovum itself, no other conclusion could be drawn, save
that this represented the impregnated ovum of the expected men-
struation. Based on these facts he estimated the age of the
ovum as possibly three to four days.
Although I agree with Peters in regard to the manstruation-
like appearance of the mucosa, I cannot coincide with him in his
calculations from the history of the case.
We read' on page 3 of Peters' work that the woman had her
last normal menstruation on September 1st, 1895. At the end
of September (on September 28th, according to the normal type,
1 Pfannenstiel 1. c. pg. 200.
2 Peters 1. c. pg. 16.
'
THE .MUCOUS MEMBRANE ELEVATION. 41
and no statement to the contrary is made) the menses should
have reappeared. They did not, however, return at the ex-
pected time, and the woman, fearing pregnancy, committed sui-
cide on October 1st. This would be 3 to 4 days after the ex-
pected appearance of the menses. Or, if we assume from certain
observations that the ovule escapes from the matured follicle
about two days before menstruation, the suicide was committed
five to six days after the expulsion of the ovum from the follicle
and the impregnated ovule presumably would then be five to
six days old. Probably it is only a lapsus calami when Peters
on page 16 states, that the woman was about to menstruate on
the day of the suicide ; he should have said that her death oc-
curred about three to four days after the expected menstruation,
which for the first time had failed to appear.
Whether or not this ovum be four or six days old does, how-
ever, not in the least diminish the value of this specimen, for
by its examination Peters has certainly succeeded in throwing
new light on the embedding theory of the human ovum.
An accurate idea of the respective age of such ova we shall
have only after more such small ova have been described, of cases
in which absolutely reliable histories have been obtained. At
present one seems justified in associating his specimen with the
first missing menstruation.
The same surely holds true of my specimen, even without a
history, since there is not even a suggestion of a embryonic
rudiment, of an amnion, etc., present, as was found .in 'Peters'
ovum.
I am, however, willing to refrain from making any estimate of
the age of my ovum. Later investigators who may have the
good fortune to work with reliable specimens of cases in which
accurate histories are obtainable, may then determine the age of
my specimen.
Of a very similar character is also the ovum demonstrated by
Graf von Spec in Kiel in 1905. It also comes from a woman
who committed suicide by taking oxalic acid. "The mucosa of
the uterine body, as is characteristic for pregnancy, was divided
into irregular areas by deep furrows. One field immediately in
front of the right tubal opening, on the anterior wall, was more
prominent and had an umbilication with a marked discoloration.
This aroused the suspicion of being the nidus of the ovum. His-
tologic sections made of this portion of the mucosa proved to be
very satisfactory for examination and on demonstration through
the epidiascope showed the following condition : Taking
up about two-thirds of the free surface of this prom-
42 THE EMBEDDING OF THE OVUM.
inent mucosa area, in a cavity of the interglandular connective
tissue of the mucosa, an ovum 1.5 x 2.5 mm. was situated. It
had but few villi and contained an embryo in a very rudimentary
stage. Between the surfaces of the chorion and uterine tissue,
here and there, small quantities of blood could be seen which es-
caped from ruptured vessels. The walls of the ovular chamber
were entirely made up of elements of the interglandular connec-
tive tissue. All the glands opened into the uterine cavity, none
into the ovum chamber. Like the ovum of Peters, the serotina,
i. e., the mucosa, between the ovum and uterine muscularis, con-
tained, within greatly dilated endothelial tubes, large blood clots
apparently formed still during life. This blood served a double
purpose, furnishing nutriment to the ovum and protecting the
deeper portions of the mucosa against the corrosive action of the
ovum. That portion of the envelope of the ovum which separ-
ates the cavum of the uterus from that of the ovum, consisted of
a thicker or thinner layer of interglandular connective tissue cov-
ered towards the uterine lumen by a single stratum of epi-
thelium."
Thus it can be seen that also in this specimen the ovular en-
velope is formed by decidual tissue, none of the uterine glands
open into it, and near the ovum, especially in the serotina, large
blood masses lie in immensely dilated endothelial tubes.
II.
THE EMBEDDING OF THE OVUM. THE CAPSULARIS.
THE FIBRIN COVER ("GEWEBSPILZ OF PETERS").
Since it has been established by Graf von Spec for the guinea
pig and by Peters for the human being, that the ovule bores its
way through the surface of the mucosa, it has become the duty
of the later investigators to determine whether this is the only
mode of nidation or whether not occasionally a circumvallation
occurs arising from the mucosa surrounding the ovum.
Pfannenstiel 1. c. page 194, is correct in saying: "The ovum de-
scribed by Peters is already completely or almost completely en-
veloped by the mucous membrane. In spite of the great value of
this specimen, in spite of the fact that it has revolutionized our
views concerning ovular nidation, we should not forget that this
is the only specimen extant of this early stage of development and
it would seem advisable not to go too far in its interpretation."
THE EMBEDDING OF THE OVUM. 43
Which, then, are the similarities and which the differences be-
tween mine and 'Peters' specimen ?
First of all, it is to be noted that the mucosa elevation on the
sides of the ovule almost up to the fibrin cover, is still covered
with a surface epithelium (Fig. 3. e. Plate II).
Close to the fibrin cover the epithelium becomes irregular, is
partly swollen and partly detached. In some, sections (Figs. 7 &
9, e,e, 'Plates IV & V) it extends like in Peters' specimen (Plate
I U.E.) under the fibrin cover, as a narrow band. At the^ slope
towards the furrow (F) it is practically absent.
Special stress is to be laid on the fact that under the center
of the fibrin cover where it lies closest to the summit of the
ovum, no more regular surface epithelium is met with. Like-
wise nowhere within the ovular chamber can even a trace of
epithelial lining be found which connects with the surface epi-
thelium.
One peculiar condition which appears only in the sections 15-19
(Figs. 5 & 6 Plate III) must be described. in detail. To the
right, below the center of the fibrin cover, a depression (e. s.)
is noticed which is lined by a few cuboidal cells (Fig. 5) whose
resemblance to epithelial cells cannot be denied. Among them,
however, are a few broader cell bodies with large nuclei which
are not unlike syncytial cells. This supposition is strengthened
by the fact that with high magnification one can see a few large
syncytial cells extending from the nearest trophoblast plugs to-
wards the depression e. s. (More details will be given in a later
description of the trophoblast, Fig. 10.)
This funnel shaped depression is still distinctly visible in Fig.
6 at e. s. Here, however, it is filled with red blood cells which
have escaped from the ovum chamber and in their- further course
cannot be differentiated from the blood cells which infiltrate the
structureless fibrin mass. (Fi.)
This funnel shaped depression seems worthy of special note
and probably deserves to be particularly considered in all fu-"
ture investigations. It is certainly the only visible and partly
preserved connection between the ovular chamber and the uter-
ine cavity.
What will be its significance? It might be regarded as the
remains of a glandular orifice, yet much speaks against this. First
of all there is no gland opening anywhere within a wide area of
this spot, especially outside of the fibrin cover. Naturally so,
since the glands have been displaced by the ovum and lie more
meridionally. We have already shown that the glands do
not take any direct part in the formation of that cavity which
44 THE EMBEDDING OF THE OVUM.
harbors the ovum and that this chamber nowhere in its interior
carries a lining of uterine epithelium. If this funnel was ac-
tually a gland orifice the good preservation of its cells would be
striking in contrast to the glandular openings still visible to the
sides of the ovum, since their epithelial cover consists almost
only of swollen, detached and liquified cells.
The question suggests' itself, however, whether this funnel
shaped depression does not indicate the site where the ovum has
entered the mucosa. My ovum does not show any "Anlage" of
an embryo and therefore must be younger than that of Peters,
in which an amniotic cavity with an exocoelum is distinctly vis-
ible. The assumption may be justified that my ovum has just ac-
complished its nidation and is about to become sufficiently an-
chored to the ovular envelope by sending out trophoblastic pro-
cesses. The fact that this place at e. s. represents the only
visible connection, obviously leads to the question, whether or
not the ovum has possibly here entered the mucosa. This place
is certainly about to become obliterated by the encroachment of
decidual cells from both sides, but there is still a relation extant
between the interior and exterior. At a place within the blood
filled space of the ovular chamber lying closest to the depression
e. s. (in Fig. 6), an escape of red blood cells can be followed, one
sees how they collect at e. s. and from here permeate the fibrinous
cover. Thus one seems justified in assuming that also the fibrin-
ous cover has originated here, that it serves more or less as a
lid and that it is supplied with red blood cells until the envelope
completely surrounds the ovum.
Before the structure and development of this fibrinous cover
is considered in detail, the process of embedding of the ovum,
and the ovular chamber must be considered. In all sections
(Figs. 14, 22 & 23, Plate VIII, XII & XIII) the decidua com-
pletely surrounding the ovum, shows the indentical histological
structure. Although varying in thickness at different places,
especially at the summit of the ovum and at both sides of the
mucosa elevation, the decidua everywhere consists of thin wavy
fibres containing larger and smaller decidua cells, numerous cap-
illaries and a large number of extravasated red blood cells. One
can therefore not say that the mucosa has reflected itself from
the two sides over the ovum, lying in its chamber, nor can as-
sert, that a scar can be seen on the summit of the ovular ele-
vation.
Everything points rather to the fact, which has been proved
by Peters, that the ovum has burrowed its way into the mucosa,
in our specimen somewhat asymmetrically in a mucosa eleva-
THE EMBEDDING OF THE OVUM. 45
tion. The point of entrance in our case has become almost com-
pletely obliterated, but there still persists a narrow channel from
which some blood oozes, the latter accumulates at the summit of
the ovum and becomes transformed into the fibrinous cover.
It Peters' specimen the connective tissue does not as yet form
so complete an envelope as in mine, in which the ovum has
possibly already penetrated to a greater depth. In his case, there-
fore, the opening still existing in the decidual envelope is covered
by a broad layer of fibrin partially organized. It is called by him
the "Gewebspilz." The trophoblastic processes with their syn-
cytial excrescences are directly attached to it
Concerning the clos.ure of the defect through which the ovum
has penetrated, 'Pfannenstiel (1. c. page 220) has expressed him-
self very carefully. Since at that time Peters' specimen was the
only one known, he considered the process of closure as uncer-
tain. Descriptions of very young ova as given by Peters, Graf von
Spec, Heukelom, Leopold and Keibel had stated "that at the point
of closure not a distinct decidual tissue but a scar tissue is found,
consisting chiefly of fibrin and showing an absence of blood ves-
sels, also he himself was able to confirm the presence of this
fibrinous scar tissue, he would not consider this question defin-
itely settled since the formation of fibrin within the decidual
envelope of the ovum is very common and often can be traced
to an entirely different cause. It would seem possible that the
final closure of the decidual capsule around the ovum is accom-
plished by a firm coalescence of the vascular connective tissue
which proliferates from opposite sides. At this place, which is
the most poorly nourished portion of the reflexa, soon after-
wards signs of degeneration would appear especially so in ova
which are more superficially embedded. The blood plug of
Peters thus might possibly be considered an abnormal condition.''
The assumption of Pfannenstiel that the ovum has an entirely
decidual envelope is confirmed by my case, with the exception
of that narrow opening at e. s. As mentioned before, it is
caused by a deeper implantation of the ovum. Nevertheless, there
is in our case, like in that of Peters, a large and very broad
fibrinous cover lying over the top of the ovum, so that such a
cover must form even in case of complete decidual inclusion. In
any explanation, however, of the origin of this fibrinous cover
one must consider this narrow opening through which the ovum
sank (e. s. Fig. 5 & 6) and whence that tiny amount of blood
has oozed to the surface and there coagulated.
In order to obtain a clear conception of this fibrinous lid one
must first of all gain a precise idea of its form, length, width and
46 THE EMBEDDING OF THE OVUM.
structure. It can be seen in the microscopic sections from 10
to 70, then again from 75 to 103 and a trace of it finally in section
119.
The entire 160 microscopic sections are embedded so that the
slope of the mucosa elevation with the furrow (F) lies to the
right of the specimen and all illustrations follow the same rule.
It will thus be perfectly plain, if in describing this fibrinous
cover the term "left," applies to the direction toward the uterine
fundus, the term "right," toward the slope of the mucosa ele-
vation and the furrow (F).
If for the purpose of reconstruction, the various sections are
placed together, one receives the impression, in a view from
above, that two minute drops of coagulated blood lie on the mu-
cosa elevation near its slope. The larger drop lies more towards
the right edge of the uterus, the smaller and flatter one to the
left. Still farther to the left in section 119 a trace of a coagu-
lated mass can be seen.
All sections through the larger drop show its right half con-
siderably thicker and more voluminous than the left. On the sec-
tions through the center, the fibrin appears in rolled up waves
like a congealing substance. (Figs. 7, 9, 11, 14, 15. Plates IV,
V, VII. VIII. IX.) In explaining this condition one must con-
sider that this thicker portion is situated exactly over the fine
fissure (e. s.) through which blood is still oozing from the
ovular chamber. On the other hand one must keep in mind that
the escaping drop, both in the lying or standing position of the
woman, must gravitate downwards, i. e., in the direction of the
slope of the mucosa elevation.
The size and structure of the fibrinous cover as well as its re-
lation to the ovular chamber can be best understood from a
study of some of the illustrations. In section 10 (Fig. 4, Plate
II) the cover suddenly appears. It has the shape of an oblong
cylinder, somewhat thicker to the right, whose left lower edge
is intimately connected with the decidual envelope of the ovum.
From the left some surface epithelial cells, already irregular,
penetrate under the edge of the cover (e). On its surface lies
a band varying in thickness that possibly consists of surface epi-
thelia of the mucosa, which, however, are not discernable as such.
The cover itself consists of a mass of fibrin in which here and
there white and red blood corpuscles are met with.
Almost identical conditions are found in sections 15 and 19
(Figs. 5 and 6 Plate III), only that in Fig. 5, in the band on the
surface, a few cuboidal epithelia can be seen, whose origin T
failed to determine.
THE EMBEDDING OF' THE OVUM. 47
In sections 22 (Fig. 7) and 27 (Fig. 8 'Plate IV) the fibrinous
cover appears in a considerably changed form. Like a catter-
pillar with a large head, it lies on the ovular envelope, which has
become thinner. Under its right and left edge (Fig. 7) traces
of surface epithelium are seen. Its superficial portion contains
a few white blood cells, in its interior a delicate network of
fibres. Most conspicuous, however, is the folded appearance of
the coagulated .mass to the right (Fig. 8 fi) which depresses the
ovular envelope.
These conditions are most plainly visible in Fig. 9 (Plate V)
and Fig. 11 (Plate VII). In the latter, representing section 30,
at fi., fibrinous masses, like frozen waves, are heaped one upon
the other, causing a deep indentation of the walls of the ovular
chamber. In Fig. 13 the fibrinous masses contain a large amount
of white blood cells, still more of them in Figs. 16 and 17, while
in Fig. 15 the clubbed end of the cover is the thickest. Here
over the depressed portion of the ovular envelope the fibrinous
masses are arranged exactly like in a corpus luteum in the stage
of organization.
Gradually the arrangement and signs of an organization make
their appearance in the fibrin cover in form of connective tissue
fibrils. Thus a sort of new formed protective layer develops
over a portion of the ovular envelope. This portion in itself is
very thin, but is further weakened by trophoblastic processes
which, as will be described later, for the purpose of fixing the
Eianlage, have penetrated into the summit of the ovular cham-
ber. (Compare Peters.)
The formation of a protective cover for which the escaping
blood droplets have been utilized, must be regarded, as has al-
ready been emphasized by Peters, an exceedingly important
process and a wise provision of nature.
If Figs. 18 and 19 (sections 68-69) are compared it will be
seen that the left portion of the fibrinous band has undergone a
certain organization and reenforces the ovular envelope. The
right clubbed shaped end is a mass of twisted fibrils which has
gradually decreased in size and like a button (Fig. 19 kn) lies
in a depression.
Since with section 70 the larger of the coagulated drops disap-
pears, it must be assumed that the surface between sections 70
and 75 (Figs. 19 and 20 Plate XI) is free of any fresh deposit
of fibrin, carries, however, the continuation of the organized
portion.
In section 75 (Fig. 20) suddenly a new fibrinous layer ap-
pears. To the left of the mass, which is in a state of organiza-
48 THE EMBEDDING OF THE OVUM.
tion, lies a band varying in thickness which resembles the rests
of surface epithelia and extends upwards along the edge of this
new fibrinous cover, thus the observation described above is prac-
tically repeated. Again the cover becomes oblong but in this
instance is thicker in the middle portion, but in general does not
attain the length or thickness of the fibrinous cover previously
described. There are a few white blood cells in its middle part.
With section 105 this fibrin band has again disappeared, here
(Fig. 24 Plate XIV) the ovular chamber being cut more later-
ally shows a thicker wall while in the preceding sections the
fibrinous cover seemed to play the role of a reenforcement for
the ovular chamber and of a protective layer, as was clearly shown
in the pictures of the first mentioned larger fibrinous mass.
Concerning a fibrinous cover (Gewebspilz) in the ovules of
Graf v. Spec (Kiel 1905) and of Beneke, we find the following
references : Van Spec says on pg. 422 : "At the site of the fun-
nel shaped depression the uterine tissue shows a defect which
must be regarded the gate through which the ovum entered the
endometrium during the process of implantation. This open-
ing is sealed by a flat blood coagulum (fibrin, containing leuco-
cytes and red blood cells). Thus the conditions are very sim-
ilar to those described by Peters. The opening which appears
with a diameter of not more than 0.8 mm., possibly is larger
than at first produced by the ovum. This increase may be due to
the stretching and* growth, possibly also to a histolytic action
of the ovular wall."
Beneke reports on page 772: "The tissue plug (Gewebs-
pfropf) which closes and fills the defect in the reflexa in general
corresponds in its histologic structure to that described by
Peters. It contains blood, fibrin, leucocytes, etc."
After this description one's attention obviously is again drawn
to the question of the similarities and discrepancies between mine
and Peters' ovum.
Up to this point both specimens are identical in all essential
features.
In both we see the decidua vera in a condition of edematous
infiltration. In both the ovule lies in a swollen mucosa near a
furrow, in both instances the ovule has penetrated into the mu-
cosa at a place deprived of epithelium and has pushed aside the
tissue and the glands so that the latter follow a meridional,
curved course. Over the top of the ovum the edges of the mu-
cosa have approached each other in an attempt to close the
ovular chamber completely. This process has progressed further
in mine than in 'Peters' specimen. In both cases on the top of
THE EMBEDDING OF THE OVUM. 49
the arch, where the ovum has entered, a blood coagulum is found
serving as a protective covering, which in Peters' specimen re-
sembles a mushroom, in mine a caterpillar with a large head.
This variation in shape so well pronounced in all my sections,
calls for a more appropriate name. Peters' term "Gewebspilz"
certainly cannot be applied to my specimen. With the idea that
such an expression should be applicable to all later cases it
seemed most natural to select the term "fibrinous cover" (Fibrin
decke). Whatever its name may be, however, the fact remains
that in these two youngest human ova it was present, large and
well developed. This explains the peculiar rests of fibrin found
on the top of the ovum in older specimens, as, e. g., described in
my atlas (Uterus und Kind, Plate I).
As first shown by Peters, our specimen also proves that the
old circumvallation theory of the formation of the reflexa is in-
correct, but still a reflexa is present. If one drops a stone into
the water the latter closes over it and if an earth worm bur-
rows itself into the ground, the hole made by it will again grad-
ually be filled with the loosened soil. 'The rapidity with which
it becomes closed will depend upon the character and structure
of the soil.
It seems permissible to call rcflc.va in Peters' and my speci-
men, those decidual arches which approach each other approxi-
mately over the center of the ovum. One must, however, keep
in mind that these arches have not newly formed and closed over
the ovum, but that as a matter of fact, they have existed from
the first beginning and that through them the minute ovule has
burrowed its way. The arches become higher the more the
ovum grows. It is obvious that the opening between the arches,
i. e., the passageway of the ovum, finally again becomes closed
by either a fibrinous plug or a fibrinous band.
In the following it will be shown that the great similarity ex-
isting between the microscopic pictures of Peters' and my speci-
men, leads to a like interpretation of the findings.
Peters (pg. 28) emphasizes that his ovum has become im-
planted in the swollen mucosa near a furrow, the same has oc-
curred in my case. Since the uterine mucosa prepared to receive
the ovum is traversed by numerous deep furrows, he thinks it
might possibly happen "that the minute human ovule could be
caught in such a groove and then actually could become em-
bedded (exactly like the ovum of the erinaceus) if the edges
of the furrow coalesced and finally closed." "In such an in-
stance, however, the ovum would be completely encircled by a
50 THE EMBEDDING OF THE OVUM.
uterine epithelium and surely one or the other gland would be
found opening into the ovum."
I agree, that an ovule in this way might enter such a furrow
and be caught, but still it could there sink into the mucosa, but
even if it should further develop while wedged in this groove,
I cannot see the necessity that it should remain surrounded by
uterine epithelium and that as Peters explains this protecting
maternal layer should then be secondarily destroyed by the pro-
liferation of the fetal trophoblast.
In the premenstrual stage the epithelium of the uterine mu-
cosa is loosened, swollen and rendered detachable by small sub-
epithelial hemorrhages to such an extent that an intact, firmly
adherent and continuous epithelial covering in such a furrow
could not be expected and hardly ever is found. Therefore,
from an anatomic histologic point of view, I cannot object
against the possibility of ovular implantation in a furrow and
would not regard it as surprising if a gland would run up to the
ovular chamber (compare Fig. 24 dr t Plate XIV) and appar-
ently open into it. Such facts do not alter my present assump-
tion that the ovum passing through the superficial epithelium and
pushing the glands aside, burrows itself into the mucosa.
In my opinion, it matters not where the ovum has become
attached and where embedded, whether on the summit of a
cotyledon (as in 'Peters' case) or near the slope of an elevation, as
in my specimen, or in a furrow, the ovule will everywhere be
the source of considerable irritation and a pronounced reaction in
the adjoining tissues. This reaction has been so well described
by Peters that I give my own opinion by simply citing his own
words (pg. 29) : "One seems justified in assuming that at the
site of implantation, either through mechanical irritation or by a
provision of nature, the tissues become highly congested, plasma
(edema) and corpuscular elements of the blood extravasate, the
preformed endothelial tubules dilate, the mucosa swells and
thickens. It still remains undecided in which manner the blood
elements are set free, whether as a result of an increased pres-
sure within the vessel or through rhexis or a free diapedesis, or
possibly the capillary vessels have been opened by the epiblast,
which has begun to proliferate as soon as it came in contact with
the maternal tissues. The fact that the fibrinous plug which lies
on the summit of the ovum pointing into the uterine lumen, con-
sists of blood elements, almost forces us to assume that during
the process of the embedding of the ovum into the edematous
connective tissue stroma, the ovum, and its free surface towards
the uterine cavity, is surmounted by a blood coagulum which pro-
THE EIANLAGE AND THE OVULAR CHAMBER (EIKAMMER.) 51
tects it and possibly furnishes nutrition for that portion of the
ovular surface which as yet has not established its connection
with the maternal tissue."
If we compare the histologic pictures of our specimen- with
this description, the sections 15 to 19 call for special attention,
because they plainly prove the minimal but quite noticeable es-
cape of blood from the ovular chamber into the fibrinous cover.
We have also seen how the left half of the fibrinous lid has be-
come more and more organized and in this way reenforced the
decidual envelope and how to the right the large head of the
fibrinous band, which lies in an indentation of the ovular cham-
ber, closes the opening of entrance.
Still other similaraties can be detected. In Peters' ovum
(Plate III, Fig. 4) there is also a lateral and central depression
in the fibrin plug, so that the fetal ectoblast almost immediately
touches it. On the other hand in his specimen the trophoblast
processes also penetrate into the decidual layers over the sum-
mit of the ovum, so that in some places they become extremely
thin and appear worm eaten.
Finally, it shall here be mentioned (Peters, pg. 44) that the
envelope of the embryo of Graf v. Spec was surmounted by a
flat blood coagulum.
III.
THE EIANLAGE AND THE OVULAR CHAMBER
(EIKAMMER).
\Vc have seen that the o.vule has burrowed itself, gradually
pushing aside the decidual tissue. Some of the glands were dis-
solved, others displaced. The blood vessels were in a state of
^narked congestion, multiplication and coalescence. This shows
clearly how the ovular chamber, especially its summit, consists
<of decidual tissue and how large blood spaces adjoin its inner
surface. These blood spaces are divided by trophoblastic col-
umns and communicate directly with the surrounding capillaries
situated in the wall of the ovular chamber.
In the next chapter these intercommunicating capillaries and
irophoblast processes with their attachment to the inner surface
oi the ovular envelope, shall be considered.
Here we shall first of all speak in general of the "Eianlage v
(Fig. 18, Plate X). An exact idea of the form of the Eianlage
52 THE EIANLAGE AND THE OVULAR CHAMBER (EIKAMMER.)
could be gained only by a reproduction of pictures of all consecu-
tive sections, which, for obvious reasons is impossible.
Studying only Fig. 18, the sections 60 to 70 and 80 and 81
in rir.te XI, Fig. 21, one gains the impression that a somewhat
irregularly shaped structure lies almost in the center of the
ovular chamber from which mesodermic processes extend from
all sides. Some of these processes are covered with an ectoder-
mal layer (Fig. 18 ekt.).
Here and there the ectoblast layer (e. g., at ek^) is detached
by an extravasation of blood under it. This does not seem
to be normal. At present, however, we must leave the ques-
tion undecided how far this condition is pathologic or still physio-
logic in this earliest stage of development. The number of very
young human ova so far known to us is still too limited, on the
other hand in our case the ovule and the entire mucosa of the
uterus is so well preserved that this one feature, even if patho-
logic, could not diminish the value of the other fiindings.
Considering the form of the Eianlage as it appears in our own
specimen, we find in a series of consecutive sections that it is at-
tached with a broad base to the inner wall of the ovular cham-
ber near its summit (Fig. 18 a). This attachment is firm and
the tissue is here also slightly permeated with red blood cells. If
permitted to make a comparison one could say that the ovule
sticks like a leech with its head to the inner wall of -the ovular
envelope, while the rest of the body is suspended in the cavity.
In order, however, to avoid any misunderstanding it is here
stated that already in this stage long trophoblastic processes ex-
tend from the irregularly shaped Eianlage (Fig. 18, tr, tr, Plate
X) which are attached to the chamber wall, and in this way, like
thin threads, fasten the apparently floating ovum.
Even under very high magnification this Eianlage is appar-
ently composed only of a congealed mass, traversed here and there
by structureless, very delicate fibres, the latter passing chiefly
along the sinuous periphery of the Eianlage. The actual contour,
however, in the main is formed by small round or oblong cells
with deeply stained nuclei which in connection with the trans-
parent network of fibres create the picture of villous tissue, i. e..
of the "Anlage" of the mesoderm.
Before entering into a consideration of the trophoblast and its
attachment, it may be well to recapitulate what has been said
before. It seems important ' to have a clear conception of the
condition of the ovum after it has entered the mucosa and of
THE EIAXLAGE AND THE OVULAR CHAMBER (EIKAMMER.) 53
the condition of the endometrium at the time when the ovum is
entering it. We assume that the most suitable time for implan-
tation of the fertilized ovum is the last few days preceding the
first missed menstruation.
In the premenstrual stage the mucosa is not only congested,
but as is shown in Fig. 25, Plate XV, permeated by extra vasated
blood. According to Gebhard, a second stage follows character-
ized by an accumulation of large extravasates in irregular lacunae
of the mucosa. As a result of the rupture of some capillaries
beneath the surface epithelium, subepithelial haematomata are
formed from which, after slight detachment of the epithelium,
blood escapes into the uterine cavity as -the visible menstrual
flow.
The blood which has escaped from the capillaries thus spreads
both beneath the epithelium and in the deeper layer of the mu-
cosa as an irregular extravasate and also enters the glands (Geb-
hard). One must therefore bear in mind that this free blood lies
chiefly in artificial cavities, not lined by an endothelium.
The fertilized ovum which in the premenstrual stage, i. e., 'be-
fore the menstrual flow has appeared, is about to enter the uter-
ine mucosa, finds a markedly loosened tissue infiltrated with blood
and will hardly experience any difficulty to pass through or push
aside the loosened surface epithelium.
When the ovum has penetrated the superficial layers of the en-
dometrium it is in a tissue permeated by the extravasates just
described and lies either free in one of these extravasates or at
least at the edge of one of them in the tissue of the mucosa. At
any rate, we seem justified in assuming, that the fertilized ovum,
immediately after its implantation, displays great vitality, that it
will develop and that above all things it will attempt to fasten
itself, which probably is most quickly accomplished by extending
the first trophoblast processes.
My idea is, that the embedded ovum, for a very short time,
either partly or completely, lies in a blood extravasate. Since
these blood spaces have formed as a result of a rupture of capil-
laries they are devoid of a continuous endothelial lining and
show an endothelium only at places where ruptured capillaries
enter.
The ovum thus floats, possibly only for the first few hours, in
a blood cavity, i. e., it is from the very beginning surrounded
by a maternal blood from which it receives its nutriment. When
and how quickly the entrance gate is closed again we do not
know, it is possible that this, process varies as to mode and time.
Many observations, however, suggest that the closure is effected
54 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
by means of a drop of blood which coagulates and later becomes
organized. (Peters, Leopold, Graf v. Spec.)
That it should be closed by a blood drop is not at all surpris-
ing, since the ovum breaking into the mucosa, so to say, has set
a small wound and has opened up a blood cavity. From the lat-
ter a small amount of blood could easily ooze. This, then, would
at the same time be the most natural process o*f restoration of
the injury produced by the process of embedding.
Without these preliminary remarks it would be rather difficult
to understand the following chapter dealing with the trophoblast.
In studying the trophoblast I had the same experience as
Peters. The microscopic pictures are so varied and oftentimes
so complex that it requires considerable time to properly interpret
the findings. One often wonders what bold conclusions were
drawn by onesself and other writers, concerning the very early
stages of development, from the study of ova representing a
much later period of pregnancy.
It is obvious that it should be the aim of both gynecologist and
pathologic anatomist to search for all available material in order
to place our knowledge, concerning the first days of embryonic
life, upon a more secure basis.
After having followed the ovule up to its entrance into that
irregular blood space of the mucosa, it becomes interesting to
understand the action and functions of the ovum for the pur-
pose of sustaining its existence. This is accomplished chiefly
by means of the trophoblast.
IV.
THE TROPHOBLAST AND ITS SURROUNDING BLOOD
SPACES. THE INTERVILLOUS CIRCULATION.
In presenting this subject the following order seems the most
appropriate : With the aid of sections 50 to 100 taken from the
center of the ovum we elucidate : 1. THE BOUNDARIES OF
THE EIANLAGE AND THE TRO'PHOBLASTIC PRO-
CESSES. 2. THE MANNER BY WHICH THESE TROPH-
OBLASTIC PLUGS BECOME FASTENED TO THE IN-
NER WALL OF THE OVULAR CHAMBER. 3. THE
BLOOD SPACES LYING BETWEEN THE TROPHO-
BLASTIC PROCESSES. 4. THE RELATION OF THESE
THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES. 55
BLOOD SPACES TO THE CONFLUENT CAPILLARIES
IN THE SURROUNDING TISSUE.
ad. 1. In Figs. 16, 17 & 18 (Plates IX & X), the ectoblast
cover and the trophoblast processes are seen. We are able to
follow both their primary condition and further development.
The ectoblast (in Fig. 18 ekt. and Ek^. Figs. 16 & 17 ekt.
and Figs. 26 & 27 Plate XVI) consists of a double row of cells.
The inner row is composed of round, in some places oval cells
adjoining each other with their pole ends and containing a dark
stained nucleus which almost fills the cell. (Langhans' cells Figs.
26 & 27 1. z.). On the outside of this cell layer we find much
larger cells which contain one or more nuclei. Many of these
cells are considerably swollen, showing a fine granulation of their
cellular substance (Syncytiurh). (Figs. 26 & 27 sy.) In fol-
lowing this ectoblastic band, which here and there is detached
from the mesoblast, in the sections 47 and 60 represented in Figs.
16 & 17, we can establish several facts. At various places (e. g.,
Fig. 17 k) by a proliferation of the Langhans' cells the inner
layer is thickened, forming small buds covered by the syncytium.
These buds gradually become thicker and longer (Figs. 16 &
17 tr & Fig. 18 tr to the right), but almost everywhere remain
covered with syncytial cells, as can be recognized in Figs. 16 &
17 under high magnification from the double layered band
marked "ekt." (Fig. 27 sy.)
The ectoblast in this stage of delevopment has already sent
out a number of short sprouts, but the mesoderm which is just
forming has not as yet entered these processes, so that one can-
not speak of villi in the usual meaning of this term.
It must, however, be emphasized that these earliest buds as
well as the further developed processes (the ectoblastic band in
Fig. 16) are completely surrounded by maternal blood and do not
come in contact nor have any connection with the maternal tis-
sue, especially decidual cells.
While this fact alone proves the close relationship of these two
cell layers to the ectoblast, furthermore, in this stage of devel-
opment of the ovum absolutely nothing speaks in favor of an as-
sumption that the ectoblast or possibly its external cover the
syncytium, etiologically has any relation to the maternal tissue.
The ectoblast is a part of the Eianlage, its buds are the expres-
sion of the development of the latter. Without an Eianlage neith-
er a formation of trophoblast nor a proliferation of syncytium is
conceivable.
Following further the increase in the size of the trophoblastic
buds (e. g., Fig. 18 tr. to the left; Fig. 16 tr to the left; Fig. 21
56 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
Plate XI; Fig. 22 tr, tr; Fig. 23 tr Plate XII & XIII) one no-
tices that the Eianlage, besides these finest processes, gradually
sends forth larger ones which tend to reach the inner surface
of the ovular chamber. Peters has very appropriately (pg. 91)
compared these trophoblastic processes to the arms of an oc-
topus.
Here also nothing can as yet be seen of an extension of the
mesoderm into these trophoblastic buds. It seems that first of
all the ovum attempts to gain a firm hold and only later it pro-
ceeds to form true villi by the proliferation of the mesoderm.
ad. 2. How and where do the ends of the trophoblastic pro-
cesses become attached to the wall of the ovular chamber?
In order to answer this question we must study not only the
sections through the middle, but especially those through the
lateral portion of the ovum. (100 to 120.) A very notable
finding can be recorded. In speaking of the Eianlage it has
been stated that it is attached like a leech with a broad head to the
summit of the ovular envelope, but that here also the attachment
is effected by a thin layer of ectoblast containing syncytial cells.
In looking over the sections through the lateral portion of the
ovum one notices that coincident with the gradual disappearance
of the Eianlage, broad trophoblastic processes and columns along
the whole periphery of the ovum but especially near its sum-
mit, like a network, connect the Eianlage with the chamber walls.
The processes and columns consist of oval Langhans' cells
darkly stained, closely pressed together and filled with round and
oblong nuclei. Everywhere, especially so laterally, they are
covered with large syncytial cells, some of them oblong irregu-
larly shaped, others standing on their edge filled with oblong
crescent shaped or with small nuclei tightly pressed together.
The ends of these processes with their syncytial cover have
penetrated deeply into the thin layer of tissue over the summit of
the ovum (Figs. 16 & 17 sy. Plate IX). Here, where the
covering fibrinous band has almost disappeared, we meet with
conglomerations of syncytial cells. Lying close together they
almost create the impression of rests of surface epithelium (Figst
12 & 13 sy. Plate VII) ; yet their connection with syncytial cells
emerging from deeper layers is so evident that any idea of their
relation to rests of epithelium must be repudiated. There is still
another fact which prohibits such an assumption. It has been
mentioned several times that from the surface of the mucosa a
narrow, irregular band resembling degenerated epithelium creeps
over the edge of the fibrinous cover. In these sections also, in
which the fibrinous cover is still missing, appearing only in later
THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES. 57
sections, this band, although somewhat uneven, again appears.
Since here and there distinct cuboidal epithelial cells are visible,
one seems justified in pronouncing it surface epithelium, it then
would be a swollen rest which has remained between the two
coagulated blood droplets lying on the summit of the ovum. For
this reason there can be no association between the epithelial like
rows of syncytium and the surface epithelium. (Compare
Peters' Plate V Figs. 10 to 13, the epithelial like arrangement of
syncytial cells.)
From this explanation one can understand how in Figs. 11 &
13 (Plate VII) syncytial cells arising from trophoblast (tr)
have become arranged in rows and finally terminate in a large
cluster of cells, (sy.)
The same holds true as mentioned before for the syncytial pro-
cess (sy.) in Fig. 10 (Plate VI), which originates from the
trophoblast.
In rather regular intervals but forming quite irregular arches
the trophoblast processes which at first have floated more or less
freely in the maternal blood spaces, approach the inner surface of
the ovular chamber (Fig. 6 tr. Plate III) and by means of
these arches subdivide the periphery of the chamber into smaller
blood lacunae. (Fig. 15 tr; Figs. 22 & 23 tr.) Thus these
lacunae develop within the trophoblastic shell of the ovum as is
so excellently shown in 'Plate I of Peters' work.
Since the trophoblast excrescences and processes, as mentioned
above, are covered with a syncytium, it becomes obvious that
these blood lacunae are lined with a syncytial layer which is
plainly visible everywhere on the concave side of the arches.
(Fig. 23 sy.) In this manner the syncytium forms a line of
demarcation between the trophoblastic framework and the blood
lacunae.
In comparing these blood lacunae in the periphery of the ovum
one notices at once a difference among them which one really
must expect and which also is noticeable in Plate I of Peters'
work. The lacunae lying near the summit are more or less
closed, while those situated in the depth and on both sides are,
by means of narrow channels, connected with the large blood
spaces in the decidua. This difference explains itself in the fol-
lowing manner: In the lacunae near the summit (Fig. 6 tr), the
closure of the arches is effected by the thin wall of reflexa or by
the fibrin cover, while this is impossible in the remaining portion
of the periphery. Here the vessel walls are opened as a result of
their rupture. (Fig. 6, c.)
Whether the lacunae appear more closed or open, in both
58 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
cases the manner in which the ends of the trophoblast processes
become fixed to them is the same. It is effected by means of
smaller and larger single syncytial cells, at times by groups of
them which glue these processes to the decidual tissue.
How intimate this attachment may become which, especially
in the tissue of the summit of the ovum, could be called an an-
choring, can be seen in Fig. 11 (Plate VII). Here the tropho-
blast, together with its adjoining syncytial cells, has buried itself
deeply into the decidual coat and advanced to "sy" with one large
giant cell. A similar condition can be observed in Fig. 11 at tr
and tr 1} where the syncytial cells radiate from the trophoblastic
processes in various directions.
In the periphery of the ovum the connecting syncytial cells at-
tach themselves to the nearest decidual trabeculae or to the re-
maining portions of the walls of the enormously dilated and rup-
tured capillaries and blood vessels. Hereby a connection of the
latter with the blood lacunae is established and maintained.
The syncytium therefore does not only serve the important
purpose of fastening and anchoring the trophoblast, but during
the further development of the ovum brings about the gradual
dissolution of more capillary walls (Fig. 28 Plate XVI) and in
this manner constantly provides larger and larger blood spaces
for the sustenance of the ovum.
Now that we have become acquainted with blood spaces which
lie between the trophoblastic processes (ad. 3), it will be neces-
sary to explain (ad. 4) the relation of these blood spaces to the
confluent capillaries in the surrounding decidual tissue. For this
purpose sections 81 to 96 and also Figs. 22 & 23 (Plates XII &
XIII) must be studied.
In looking over these sections one must keep in mind the
fact that the ovum has entered a tissue which as a result of the
premenstrual state was highly congested. Blood vessels as well
as glands were greatly dilated. Blood extravasated from the cap-
illaries, passed into the interstitial tissue and finally also found
its way into the glands. Thus the Kianlage was surrounded and
the ovular chamber formed by a very loose and spongy tissue
infiltrated with blood consisting in the main of dilated capil-
laries, many of them ruptured.
No matter how many specimens we examine and compare, the
same condition will be noticed in all. The inner wall of the
ovular chamber, as already mentioned, is formed partly by di-
lated capillaries and partly by open blood spaces with their inter-
vening framework. Since the open blood spaces only recently
have been closed capillaries, their inner wall, not everywhere but
THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES. 59
almost throughout, is lined with an enclothelium. In the im-
mediate neighborhood of the blood lacunae, in the loose tissue,
numerous large syncytial cells are found singly or in clusters,
some of them lying close to the chamber wall. These cells, so to
say, are the pathfinders for the syncytimn, which glues the
trophoblastic processes to the wall. For this reason a great num-
ber of large syncytial cells are met with in the sections which
pass far laterally through the ovum. (Fig. 4 sy. Plate II.)
It is noteworthy that wherever a syncytial cell lies close to
a vessel wall the corresponding endothelial cell on the inside, in
comparison to others, seems enlarged and contains a very large
nucleus which either is oblong or undergoing segmentation.
One could feel inclined to think that the transformation of the
endothelium into syncytium is the primary result and that the
syncytium develops further from here. One then could con-
clude that the syncytium in the main originates from vessel en-
dothelium. In our specimen, however, no support whatever can
be detected for such an assumption. As a matter of fact, the
conditions are as follows : The syncytial cells which cover the
trophoblastic processes and attach themselves to the walls enter
the tissue, become disseminated between the capillaries and
glands especially in the remaining framework of tissue. They
attach themselves to the walls of the capillaries which are still
closed, corrode them and thus constantly open new blood
spaces. 1
This process, e. g., is represented in the sections 86 to 89. Here
we see large rows of syncytial cells which advance from a
trophoblast process towards the wall of a transversely cut cap-
illary and attach themselves to cells of the wall. (Fig. 28 sy.
Plate XVI.) In the next section syncytial cells pass into a ves-
sel wall. Its various layers become detached from each other.
The tissue cells which formerly lay in closed arches as well as
the endothelial cells are now unraveled. In the next sections we
see blood cells oozing from these unraveled vessel walls. The
syncytial cells force their way deeper and deeper into the lumen.
Finally one sees only half of the wall of the vessel until this, too,
is dissolved. In this way the peripheral blood spaces, one after
the other, are destroyed by the never ceasing action of the syn-
cytial cells and from these findings the conclusion must be drawn
that already, /. c., even in this early stage of development, the
intervillous spaces are indicated and even existing.
1 At the Congress in Kiel, Peters demonstrated drawings which show beautifully how
endothelial cells become detached and necrotic exactly at those places where syncytial
cells have attached themselves to the outside of capillaries; another proof that endothelial
cells do not transform into syncvtium.
60 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
Correctly we cannot as yet speak of intervillous spaces be-
cause the mesoderm has not yet entered trophoblastic process so
that villi in the strict sense of the word do not exist. In Peters'
specimen (Plate II) they have already formed. Therefore, it
would be more appropriate in our ovum to apply to these blood
spaces the term "intertrophoblastic."
It seems, however, not desirable to add new difficulties by in-
troducing a new term. The reader acquainted with the subject
will know what is meant and it would seem permissible to speak
of an "Anlage of intervillous spaces" even in this stage, since
very soon villi actually will enter these trophoblastic excres-
cences.
What can be attributed to the findings?
I am well aware of the fact, that although it lies completely in
situ and has been carefully handled, still it shows in a few sec-
tions, an excessive amount of blood which, here and there, has
pressed the Eianlage together and detached the ectoblast. Ow-
ing to this I hesitated in publishing my findings, but since in the
main all those conditions which I wish to demonstrate were so
plainly visible, and so instructive, I was finally persuaded to do
so by 'Prof. Graf v. Spec.
I believe that the description of my ovum is of value so long as
no other ovum of the same age or possibly a younger one still
better preserved is known. I will therefore compare my find-
ings with those made on the youngest ovum known in literature.
Peters has extensively quoted all those writers who have de-
scribed older ova and carefully compared his findings. Thus I
am justified in limiting myself here to his description and his
critical review of the literature on the subject, and shall empha-
size only points of importance in these new findings. For a con-
sideration of this subject, only a few of the recent contributions
will be considered, and this will also appiy to the following chap-
ter dealing with the syncytium : Pfannenstiel in Winckels Hand-
buch der Geburtshilfe, Rossi Doria, 1 Marchand, 2 Friolet, 3 Herr-
mann and Stolper, 4 Webster, 5 Graf v. Spec and Beneke 1. c.
Peters' description of the trophoblast corresponds in all es-
sential features with ours. He found the development, the
1 Ueber die Einbettung des menschlichen Fies. studiert an einem kleinen Ei der zweiten
Woche. Arch. f. Gynaekologie 76.
2 Beobachtungen an jungen menschlichen Eiern. Anat. Hefte. 21. B.
3 Beitrag zum Studium der menschlichen Placentation. Leipzig, Thieme. 1904.
4 Zur Syncytiogenese beim Meerschweinehen. Wien, Hoelder. 1905.
5 Die Placentation beim Menschen. Uebers. von Kolischer. Berlin, O. Coblenz, 1906.
THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES. 61
structure and the lining of the trophoblast processes with syn-
cytium exactly the same as I saw them. He says on page 49:
"From the simple cuboidal cells of the central portion of the
trophoblast one can successively follow, towards the surface, their
transition into the large syncytial formations/'
"Those blood lacunae which on their proximal surface carry a
layer of ectoblast cells, are almost completely lined by a thin
layer of protoplasm in which cell membranes not discernible and
in which nuclei are visible, arranged in rows. In transverse sec-
tions these nuclei appear spindle shaped, in oblique and longi-
tudinal sections they look flat, often irregularly bent and in their
appearance seem identical with the nuclei previously described
in the peripheral layer of the ectoblast. This thin protoplasma
coat which in the central portion of the blood lacunae resembles
a layer of epithelium, continues on the lateral walls of the lacunae,
thus enclosing the trophoblast columns."
Peters therefore also finds the blood within the lacunae sep-
arated from the trophoblastic excrescences and columns, espe-
cially within the arches, by a syncytial lining.
As far as that area of the decidua compacta is concerned,
which immediately adjoins the trophoblast (Umlagerungszone
of Peters pg. 52), here surrounding the entire ovum, especially
on the side of the serotina, greatly dilated maternal blood vessels
and a large number of transverse sections through glands are
seen,. The uterine mucosa here is not only edematous, but also
abundantly infiltrated with blood elements.
Further away from the ovum in Peters' specimen (Plate I) a
large endothelial tube is visible from which a number of larger
and smaller vessels branch off. These encircle the ovum like
meridians the globe. "They communicate with the blood lacunae
of the trophoblast near the oyular pole, often only after having
first passed through a part of the 'Umlagerungszone' or through
intermediate layers lying between it and. the compacta. In other
places these vessels after only a short course open into the la-
cunae at points near the serotina."
As regards the relation of the trophoblast to the surrounding
capillaries and their endothelial tubes, Peters' investigations (pg.
59) show that the trophoblastic processes do not bore their way
into the endothelial walls as I have described, but that the blood
has actively broken into the trophoblast. This condition Peters
has depicted in that excellent illustration, Fig. 23, v Plate A, to
which I cannot furnish an equal from any one of my sections.
I have only been able to observe, as stated above, that at first the
ovule lies in a premenstrual extravasate of blood in the mucosa,
62 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
that next the trophoblast grows, by means of syncytial cells,
which it sends out into the surrounding tissue, corrodes the next
capillary and thus brings one capillary after the other in com-
munication with the blood lacunae. Since the great vascularity,
however, in the neighborhood of my ovum possibly is abnormal,
I am willing to admit that most probably Peters' histologic find-
ings are more correct and therefore his deductions more reliable.
It must be added, however, that this difference in the findings is
of little importance, since I could show exactly like Peters a
point of much greater importance that a communication exists
between the blood spaces within the trophoblastic arches and
those lying farther away. We established in this way the fact,
that at this early time intervillous blood spaces exist in a rudi-
mentary and fully developed state.
Mention must be made of a rather important finding by Peters
on an ovum 3x5 mm. large (i. e., much larger than my ovum),
obtained from a woman who died of phosphorus poisoning.
"With due regard to the fact that this was a case of phosphorus
poisoning he states that in this ovum in which the trophoblast
layer was already reduced to the double layered chorion epithelium
(Langhans' cells plus syncytium) and which with its villi was
embedded in a lake of blood, the intervillous space by means of
a continuous layer of fibrin which completely surrounded the
ovum, was separated from the compacta, pathologically changed
by a suffusion of blood. Whether this fibrin layer by compari-
son with undoubtedly normal specimens can be regarded as the
primary stage of the fibrin layer of Nitabuch is a question which
cannot be answered."
I must mention that also, in my ovum, in some sections, espe-
cially near the border of the greatly dilated and open blood spaces
which surround the ovum like a shell, such strips of fibrin are
visible. I should like to refrain, however, from positively defin-
ing their significance.
The following points which have been clearly established by
the descriptions given above must be emphasized : The formation
of the blood lucanae in the trophoblast does not stand in any di-
rect relation to the destruction and dissolution of the glands, a
point which also has been made by Peters, (pgs. 75 & 76).
Nowhere is syncytium formed from glandular epithelium.
I cannot agree with Pfannenstiel, who, also in all essential
points in accord with Peters, holds different views in certain
questions, e. g., concerning the mucosa surrounding the ovum.
His observations are made on a comparatively too old an ovum,
being almost two weeks old. He asserts (pg. 242) that in his
THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPAOES. 63
belief the blood lacunae in the trophoblast are new formed cap-
illaries of the decidua, whose walls either the endothelium or
the surrounding connective tissue had transformed into syncy-
tium. In none of my sections could I find any support of this
view.
Like Peters, I found that the partitioned lacunae at the bor-
der, for the very reason that they are lying between the tropho-
blast columns and in fact are formed by them, are lined with a
layer of syncytium which ceases where the ends of the tropho-
blastic processes peripherally reach either the framework of the
mucosa or the walls of the open blood vessels. It seems ques-
tionable to me whether these lacunae, although of maternal or-
igin, can be called new formed capillaries of the decidua. This is,
however, a point of minor consequence. Of much more import-
ance is the fact, that the syncytial coat of these lacunae, as has
been demonstrated above, derives its existence from the Eianlage,
i. e., from the ectoblast.
Most noteworthy findings have been made by Rossi Doria, who
examined an ovum dating from the beginning of the second week
of pregnancy. Aside from many observations which are fully
in accord with those made by Peters and myself, his findings con-
cerning the trophoblastic capsule are especially interesting.
He also claims that the fertilized ovum penetrates the ex-
tremely congested mucosa of the premenstrual stage and enters
a space filled with extravasated blood. "The chorionic tropho-
blast excrescences from the ectoderm proliferate through the
blood until they reach the surrounding connective tissue. They
spread out with it and cause certain changes." These consist in a
dilation and rupture of capillaries and in the formation of new
extravasates in which the trophoblast forms new and longer pro-
cesses. A formation of new blood vessels which has been claimed
by Pfannenstiel has not been observed by Rossi Doria.
According to Rossi Doria, "the trophoblast corrodes the ves-
sel walls and penetrates them with the syncytium. In defense the
maternal tissue protects itself against the invasion of the tropho-
blast by means of a barrier of decidual cells (Umlagerungszone).
In this manner both the fetal trophoblastic and the maternal cap-
sules are formed."
This quotation alone proves sufficiently the great similarity be-
tween. Rossi Doria's and my descriptions, yet it may not be amiss
to also point out the harmony of our views concerning the forma-
tion and lining of the trophoblastic lacunae. I will cite verbatim
his description (pg. 503) since my observations are identical.
"The syncytium breaks into the decidual coat especially into
64 THE TROPHOBLAST AND ITS SURROUNDING BLOOD SPACES.
the perivascular spaces and beneath the endothelium as well as
into the lumen of the blood vessels. All the giant cells which
have been found by some observers in the decidua, in this stage
of development of the ovum, are to be considered cells derived
from the syncytium. (Compare my identical findings in Fig. 4).
They have become separated from a pedicle at first existing and
then even in a series of sections show the character of isolated
cells. Often, however, they remain attached to the syncytium
from which they originate, and then only by mistake can be pro-
nounced giant cells."
According to Marchand (pg. 262) the developing ovum reacts
upon the surrounding uterine inucosa like a malignant growth,
it "eats its way" into the mucosa as Graf v. Spec has demon-
strated for the earliest stage of the ovum of the guinea pig.
Other claims of Marchand need not be considered here since
the ova described by him are much older, two of them being
in quite a defective condition.
The ovum described by Friolet and estimated to be from three
to four weeks old also seems of limited value in a consideration
of the very early stage. He agrees in all main points with 'Peters.
Herrmann and Stolper have made exhaustive researches con-
cerning the ovum of the guinea pig and have arrived at the fol-
lowing conclusions: "1. In the guinea pig there exists but one
kind of syncytium and this is positively fetal, having developed
from the Anlage of the placenta. 2. There exists definite rela-
tions between the syncytium and maternal vessels ; and, 3. The
villus of the placenta of the guinea pig in a certain stage of de-
velopment, exactly like in the human placenta, shows the char-
acteristic double layer of epithelial cover." Thus the similarity
between mine and their findings is obvious.
Concerning the very meritorious work of Webster, it must be
stated that in the main it presents a clear survey of all the findings
which have been made in all the various months of pregnancy,
but does not deal with any personal investigation of a new young
ovum.
Beneke, who has examined a considerably older ovum (4.2x
2.2x1.2 mm.) with an embryo 1.86 mm. long, comes to deduc-
tions which are identical with Peters' and those of mine. In the
main he confirms the views of Van Heukelom, Peters, Marchand
and others concerning the structure of the trophoblast. He also
considers the syncytial giant cells as originating from fetal ecto-
blast. He did not succeed in differentiating a Symplasma glandu-
lare, conjunctivum and cndothclialc from fetal syncytium as has
been done by Bonnet. The synctium displaces the endothelial
THE SYNCYTIUM. 65
cells of the dilated decidual blood vessels and the epithelium of
the glands. A detailed report of his findings is still outstanding.
V.
THE SYNCYTIUM.
In the previous chapters, in describing the ectoblast and tropho-
blast, mention has so frequently been made of the syncytium
that no doubt can be left concerning my own views of its origin.
Since the opinions of most prominent investigators- are still
at variance on this point, I shall once more consider the origin
and function of the syncytium and shall quote disparaging views
of others.
In accordance with Peters, my specimens (Figs. 16 and 17)
show that the syncytium forms the external cell layer of the
ectoblast and that it is first noticed in this situation. We have
further observed that when the inner layer of cells (Langhans'
cells) begin to send out small excrescences (Fig. 17 kn.) or
longer processes, (Figs. 16 and 17 tr) these are covered with
syncytium which closely follows all the larger and largest tropho-
blast columns and with them reaches the connective tissue of the
ovular chamber.
In this manner the syncytium separates all the proliferations
of the trophoblast from the surrounding blood.
Next it becomes the duty of the syncytium to connect the ends
of the trophoblastic processes with the adjoining tissue, to corrode
the surrounding vessels, to unravel their walls and finally to
open up more and more blood spaces from which the growing
ovum may derive its nutrition.
. In all specimens thus the syncytium can be followed from with-
in outwards, from the ectoblast towards the ends of the tropho-
blastic processes, but not in the reverse direction. It can be
observed at the ends of the short trophoblastic excrescences,
which have not as yet become adherent, in form of those well
known processes which vary in shape but as a rule are knob-
like. In studying these pictures and comparing them with others
it seems impossible to come to any .other conclusion than that
the syncytium is solely of fetal origin and that it forms the ex-
ternal cell layer of the ectoblast providing for the growth and
nutrition of the ovum.
The manner also in which the syncytial cells corrode, weaken
and dissolve the vessel walls, leads us to but one conclusion, viz :
that they must be fetal tissue originating in the Eianlage. One
can see how thev leave the ovum and advance into the tissue
66 THE SYNCYTIUM.
i
but never take the opposite direction. If the latter were true,
a far larger number of syncytial cells would be found in the
immediate neighborhood of the vessels of the ovular envelope.
The fact that all trophoblastic processes, short and long, carry
syncytium and that the arches of the trophoblastic lacunae are
almost completely lined with a syncytial coat can hardly be
brought into harmony with Pfannenstiel's claim, that the vessel
endothelium is transformed into syncytium and that starting from
here the syncytium gradually proceeds toward the ovum.
Peters, with whom I agree, expresses the following precise
views concerning the genesis, the early appearance, and the loca-
tion of the syncytium (pg. 87.) : "We occasionally, though rarely
find within the trophblast, blood lacunae which as yet are not
lined with syncytium. Otherwise, however, the syncytium is
found distributed over the entire surface of the ovum, i. e., in all
sections of the complete series here and there we find bands and
masses of syncytial protoplasm varying in size, which either lie
flat on the trophoblast or are connected with it by a irregular net-
work of protoplasm or lie free in the blood lacunae or at times
push forward toward the Umlagerungszone." On these
facts Peters declares the syncytium to be a tissue of fetal origin
and my specimens have forced me to the same conclusion.
This seems to settle the question whether the syncytium could
possibly originate from uterine epithelium or decidual tissue.
Peters has advanced so many and convincing arguments in oppos-
ing this view that I feel unable to add new ones.
The following are some of the points in which my views have
changed as a result of the study of this youngest ovum and are
now at variance with views formerly expressed : The origin of
the Langhans' cells, the decidua of uterine epithelium in the
neighborhood of the ovum (already corrected on a previous occa-
sion), the origin of the intervillous spaces, and a few others.
A further advance of our knowledge of this problem depends
upon the discovery of more very young human ova in situ, com-
ing if possible from patients who have died suddenly but not as
suicides. These specimens must be carefully preserved, pains-
takingly examined and the microscopic pictures reproduced in
illustrations which are true to nature.
EXPLANATION OF FIGURES.
Plate I. Fig. 1. The pregnant uterus split up its anterior wall. The light spot
marked "ovum" denotes the site of embedding of the ovum. o. i.=
Internal Os.
Fig. 2. Three microscopic sections demonstrating the mucous membrane
elevation (h), in which the ovum lies, g line of separation of
mucosa and musctilaris. 2a Section 5; 2b~ Section 80; 2c
= Section 160. (Natural size.)
Plate II. Fig. 3. Section 4. Outer border of the mucous membrane elevation, e =
surface epithelium, v = decidua vera. d Dilated and folded gland,
showing beautifully preserved epithelium, c Capillaries. F
Mucous membrane furrow. Obj. A A. Oc. 3.
Fig. 4. Section 10. fi = Fibrin cover, e = Rests of surface epithelium, c
= Confluent blood vessels. F = Furrow, tr = Trophoblast pro-
cesses, sy = Syncytial cells. Obj. A A. Oc. 3.
Plate III. Fig. 5. Section 15. Obj. AA. Oc. 12.
fi = Fibrin cover.
w = Rests of cubodial epithelium on the fibrin cover.
e = Underlying surface epithelium (?).
e.s. = The same depressed, simulating a gland opening. (See
Text!).
c = Confluent blood spaces.
Fig. 6. Section 19. Obj. A A. Comp. Oc. 4.
fi = Fibrin cover.
e.s. Trace of depression as seen more plainly in Fig. 5.
(Section 15).
c = Confluent blood spaces.
tr = Trophoblast processes.
F Mucous membrane furrow. Red blood corpuscles between
the depression, (e. s.) and fibrin cover.
Plate IV. Fig. 7. Section 22. .Obj. A A. Oc. 3.
fi = Fibrin cover.
e From the edges surface epithelium ( ?) is seen passing
beneath the fibrin cover. Possibly syncytium.
d.v.= Decidua vera.
c = Confluent blood vessels.
tr = Trophoblast processes.
F = Mucous membrane furrow.
"ovum" = The Fianlage, laterally cut, is seen for the first time.
Fig. 8. Section 27. Obj. A A. Comp. Oc. 4."
fi = Fibrin cover. To the right arranged in folds.
d.v.^ Decidua vera.
c Confluent blood vessels.
F = Mucous membrane furrow.
dr = Glands.
tr Several trophoblast processes have become anchored in
the periphery of the mucous membrane.
sy = A plug with knob-like syncytial processes.
"ovum".= Eianlage becomes more distinct.
Plate V. Fig. 9. Section 26-28. Zeiss Obj. A A. Oc. 3.
fi = Fibrin cover. To the right arranged in folds.
e = Border epithelium (?), passing from surface under fibrin
cover. Probably a band of syncytium passing from tropho-
blast (tr).
tr = Trophoblast processes.
67
68 EXPLANATION OF FIGURES.
Plate VI. Fig. 10. The left half of Fig. 9 highly magnified. Obj. Zeiss. Aprochr.
4.0 mm. Oc. 6.
fi = Fibrin cover.
e = Border epithelium.
tr = The trophoblast processes are connected with the syncy-
tial cells, which might be mistaken for surface epithelium. (Com-
pare Peters' Plate V. Fig. 12 and 13.)
Plate VII. Fig. 11. Section 30, Obj. AA. Oc. 8. The middle part of the Section was
purposely omitted.
fi = Fibrin cover, symmetrical to the left, to the right many
folds present, as in a corpus luteum.
tr = Trophoblast processes terminating in a
sy =syncytial club shaped ending.
e = Epithelial rests (?). Probably a band of syncytium com-
ing from the trophoblast (tr).
Fig. 12. Section 32. Obj. Apochr. 4,0 mm. Oc. 3.
tr = Trophoblast processes.
sy = Syncytium.
Fig. 13. Section 39. Apochr. Obj. 4,0 mm. Oc. 3.
fi Fibrin cover.
sy Syncytial bands, terminating to the right in a club-shaped
mass of cells,
tr = Trophoblast covered with (sy) syncytium.
Plate VIII. Fig. 14. Section 37. Obj. A A. Oc. 3.
fi Fibrin cover.
e = Rests of surface epithelium (?).
c = Confluent blood vessels.
F = Mucous membrane furrow.
dr = Glands. The lateral ones (to the left and above) open
into the cavum uteri, to the right and below, into the furrow.
The middle gland (m) divides into two branches.
The right branch approaches the ovular chamber, but does not
open into it.
tr = Trophoblast processes.
sy = Mass of syncytium.
Plate IX. Fig. 15. Section 40. Obj. A A. Oc. 8.
fi = The club-like end of the fibrin cover showing the folded
arrangement as in a corpus luteum.
tr = The arch-like arranged trophoblast can be traced to the
epithelial-like bands beneath the club-like end of the. fibrin cover.
Question, whether epithelial rest or not.
Fig. 16. Section 47. Obj. A A. Oc. 8.
fi = Fibrin cover partly undergoing organization.
ei Eianlage.
ekt = Ectoderm.
tr = Trophoblast processes.
sy Syncytial buds.
Fig. 17. Section 60. "Obj. A A. Oc. 8. Lettering as in Fig. 16.
k Trophoblast buds with a syncytial covering. (See Fig. 16.
under high magnification).
Plate X. Fig. 18. Section 63. Obj. D. Oc. 3.
fi.o.= Fibrin covering undergoing organization,
ei = Eianlage.
ekt and EMi~= Ectoderm,
tr = Trophoblast processes.
a = Eianlage lying with its broad surface to the inner wall of
the summit of the ovum.
Plate XI. Fig. 19. Section 69. Obj. AA. Oc. 3.
Kn^ The fibrin cover presents itself in its right process as
a small structureless elevation.
EXPLANATION OF FIGURES.
69
Fig. 20.
Fig. 21.
Plate XII. Fig. 22.
Plate XIII. Fig. 23.
Plate XIV. Fig. 24.
Plate XV. Fig. 25.
Plate XVI. Fig. 26.
Fig. 27.
Fig. 28.
Section 75. Obj. AA. Oc. 3.
Kn The knob-like projection again becomes longer and lies
upon the organized fibrin.
s.b. = A cloddy band covering same.
Section 80. Obj. A A. Oc. 4.
fi = The structureless fibrin cover again becomes longer and
is similar to sections 10-25.
s.b. = A cloddy band lying upon same. Eianlage with numerous
trophoblast processes. (Sketched).
Section 92. Obj. AA. Oc. 3.
fi = The fibrin cover partly covered by cloddy band, (s. b.)
again becomes longer and lies upon the ovular summit.
F = Mucous membrane furrow.
c. ci, cs= Confluent dilated blood spaces.
dr ' Elongated glands surround the ovum.
tr = The trophoblast processes are anchored in the border
of the ovular chamber by means of the syncytium.
Section 94. Obj. AA. Oc. 6. f
fi = The fibrin cover with its short cloddy band (s. b.), has
again become somewhat longer and lies closely applied to the
border tissue.
F = Mucous membrane furrow.
c, ci, c-= Confluent dilated blood spaces.
dr. dn. dr, = Glands. Those marked dn and drc in the
illustration, reach almost to the uterine cavity. They are partly
filled with blood and toward the base of the ovular chamber they
appear partly displaced and partly dissolved. No gland is seen
opening into the ovular chamber.
tr = Several trophoblast processes are anchored to the inner
u-;:ll of the ovular chamber.
Section 105. Obj. A A. Oc. 8.
The fibrin cover is no Jonger present.
F = Mucous membrane furrow.
c = Confluent blood spaces.
dr. dn Glands. The one marked dn reaches almost to the
ovular chamber.
tr = A number of trophoblast processes anchor themselves to
the inner wall of the ovular chamber partly with syncytial off-
shoots.
Section 142. Obj. AA. Oc. 3.
c = Mucous membrane elevation with innumerable confluent
blood spaces.
dr = Four large glands showing a partly tortous course with
opening at dr.m.
bl = Blood free in the tissue.
Section 52. Obj. 4,0 mm. Oc. 3.
Ectoderm highly magnified (See Figs. 16, 17 and 18).
1. z. =Langhans' cells on the inside.
sy = Syncytial cover on the outside.
Section 60 (See Fig. 17) under low power.
Ectoderm under high magnification.
1. z. = Langhans' cells forming trophoblast cells and offshoots.
They are covered externally like ectoderm bands in Fig. 26 with
syncytium. Obj. 4,00 mm. Oc. 3.
Section 90. Obj. 4,0 mm. Oc. 3.
sy = The syncytial cells advance from the trophoblast toward
the wall of a transversely cut capiliary (ca) and corrode the vessel
wall (en).
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