180 THE DEVELOPMENT OF THE CHICK

of the anterior intestinal portal becomes incorporated in the

floor of the intestine as the anterior intestinal portal retreats
backwards, and the original dorsal liver diverticulum therefore
becomes anterior or cephalic and the ventral becomes posterior
or caudal (Fig. 103 B). Before this transposition occurs, how-
ever, the diverticula have grown forward towards the sinus
venosus in the ventral mesentery of the stomach, the anterior
diverticulum above and the posterior diverticulum below the
ductus venosus. The stretch of entoderm between the two liver
diverticula thus lies in the angle made by the union of the two
omphalomesenteric veins. At the stage of 26 somites, the anterior
diverticulum has grown forward above the ductus venosus
to the level of the Cuvierian veins and is large and flattened
laterally. The posterior diverticulum is barely indicated at this
time.

The anterior diverticulum was originally described as left and the

posterior as right (Goette, 1867), and this description was taken up
by Foster and Balfour. This was corrected by Felix (1892). Subse-
quent writers do not agree exactly as to the time or precise relations
of the diverticula; however, it is generally agreed that the two diver-
ticula are subdivisions of a common hepatic furrow, inasmuch as the
entoderm between them lies below the level of the entoderm in front
and behind (Fig. 103 B). Brouha maintains that at first the hepatic
furrow lies in front of the anterior intestinal portal, and that the latter
secondarily moves forward so as to include the hepatic furrow, which
later again comes into the floor of the intestine with the definitive retreat
of the anterior intestinal portal. This view does not rest on very secure
evidence, and probably based on interpretation of slight individual
is

variations as successive stages of development. Choronschitzky places the

time of appearance of the hepatic diverticula at about the thirty-sixth
hour. It is probable, however, that this is too early. I have found the

unmistakable diverticulum at a stage of 22 somites, a slight rudi-

first

ment of the anterior diverticulum in the anterior intestinal portal.

At the 30 s stage the anterior or dorsal diverticulum has ex-

panded much more, mainly to the left of the middle line, as though
to embrace the ductus venosus, and the posterior or ventral
diverticulum has an even greater development and embraces
the right side of the ductus venosus, but it does not extend as
far forward as the anterior diverticulum. Both diverticula
now branch rapidly and profusely, forming secondary anasto-
 FROM TWELVE TO THIRTY-SIX SOMITES 181

moses where branches meet, so that a complete ring of anas-

tomosing columns of hepatic cylinders is rapidly formed around
the center of the ductus venosus
(Figs. 103 B and 104, cf. also Figs.

119 and 120). But the anterior

and posterior ends of the ductus
venosus are not yet completely
surrounded by the basket-work of
liver substance, owing to the ab-
sence of any part of the posterior
diverticulum in its anterior por-
tion, and of the anterior divertic-
ulum in its posterior portion.
The floor of the intestine be-
tween the anterior and posterior
liver diverticula is depressed; later
it becomes separated from the
intestinal cavity to form a tem-
porary common bile-duct; which
then receives the two primary di-
verticula (Figs. 103 B, 104 and
187).
The pancreas arises from a dor-
saland a pair of ventral primorclia.
The former is an outgrowth of
the dorsal wall of the intestine
immediately above the posterior
liver diverticulum (Figs. 103 B
and 104). At the 35 s stage it is

a solid thickening of the dorsal

Fig. 104. — Reconstruction of the
liver of the chick at the end of
wall of the intestine of consider-
the fourth day of incubation.
able extent; a little later the base (AfterHammar.)
of the thickening is hollowed out, du., Duodenum. L., Substance
of liver. Other ahbreviations as
and the free margin sends ofT solid
before.
buds into the dorsal mesentery
just behind the stomach. The ventral primordia arise from the
posterior liver diverticulum in a manner to be described later
(Chap. X).
Mid-gut. At the 35 s stage the mid-gut is still open to the
yolk-sac. Its subsequent history is given in Chapter X.
182 THE DEVELOPMENT OF THE CHICK

Anal Plate, Hind-gut, Post-anal Gut. and Allantois. At about

the 14 s stage a thickening of the ectoderm in the middle line
just behind the primitive streak extends towards the entoderm
which is folded up so as to neariy meet it, thus cutting off the
extra-embryonic mesoblast from the primitive streak. The ecto-
derm and the entoderm then come into contact here, and form
a firm union, the anal plate (Fig. 70), which is subsequently
perforated to form the anus. At first, however, the anal plate lies
entirely behind the embryo, and the post-anal portion of the
embryo arises from the thickened remnant of the primitive streak
(tail-bud) which grows backwards over the blastoderm beyond
the anal plate. Even before this, however, the hind-gut begins
to be formed by a fold of the splanchnopleure directed forwards
beneath the tail-bud, and the hind end of the tube thus formed
ends at the anal plate (Fig. 70). The entoderm in front of the
anal tube is fused with the substance of the tail-bud, and as the
latter grows backwards beyond the anal plate it carries with it
a pocket of the hind-gut, and this forms the post-anal gut (Fig.
80).
The formation of the tail brings the anal plate on to the ven-
tral surface of the embryo at the junction of tail and trunk, and
the post-anal gut then appears as a broad continuation of the
hind-gut extending behind the anal plate, and ending in the tail
at the hind end of the notochord (Fig. 80). The further elonga-
tion of the tail draws out the post-anal gut into a narrow tube
lying beneath the notochord in the substance of the tail; it
then gradually disappears and leaves no trace.
The formation of the hind-gut takes place prior to the for-
mation of the embryonic body-cavity at this place. It thus
happens that the splanchnic mesoderm, forming the floor of the
hind-gut, is directly continuous with the somatic mesoderm.
When the body-cavity does penetrate this region it is without
direct lateral connections with the extra-embryonic body-cavity,
so that the connection of the splanchnic and somatic mesoderm
persists,forming the ventral mesentery of the hind-gut (Fig. 81).
This is a thick mass of mesoblast binding the hind-gut to the
somatopleure. The hind-gut is deep from the first, and its ven-
tral division soon begins to extend into the ventral mesentery

as a broad evagination, the allantois (see p. 143).

 FROM TWELVE TO THIRTY-SIX SOMITES 183

VI. History of the Mesoderm

The history of the extra-embryonic mesoderm is considered
sufficiently in the first part of this chapter. The history of the
embryonic mesoderm be considered under the following
will

heads: (1) Somites, (2) Intermediate Cell-mass, (3) Vascular

System, (4) Lateral Plate and Body-Cavity, (5) Mesoblast of
the Head.

0tMj^jff§^3^:

Fig. iUo. — Embryo of aboul 27 somites drawn in alcohol by re-

flected light; upper side, x 10.
Am., Amnion, ot., Otocyst. t. F. Am., Tail fold of amnion.

(1) Somites.The rate of formation of the somites from the

segmental plate and their number at different times is given in
the normal table of embryos (p. 68), and may be seen in various
184 THE DEVELOPMENT OF THE CHICK

figures of entire embryos. The formation of new somites con-

tinues after the end of the period discussed in this chapter,
up about the sixth day.
to Each somite has a definite value in
the developmental history.

Fig. 106. — The same embryo from beneath, x 10.

a. i. Anterior intestinal portal.
p., A. V., Vitelline artery.
Int., Intestinal groove.

In an embryo of 42 somites (about ninety-six hours), the value of

the somites as determined by their relations and subsequent history
is as follows:

1 to 4. Cephalic entering into the composition of the occipital region

;

of the skull.
5 to 16. Prebrachial; i.e., entering into the region between the wing
and the skull.
17 to 19. Brachial.
20 to 25. Between wing and leg.
 FROM TWELVE TO THIRTY-SIX SOMITES 185

26 to 32. Leg somites.

33 to 35. Region of cloaca.

36 to 42. Caudal.
More somites are formed later, the maximum number recorded being
52 (see Keibel and Abraham, Xormaltafeln). In an eight-day chick
the number of somites is again about 42, including the four fused with
the skull. Thus the ten somites formed last are again lost. This points
towards a long-tailed ancestry for birds.

Each somite is composed of an epithelial wall of high, columnar

cells, enclosing a core of cells that nearly fills the cavity (Figs.
112, 113, etc.). From each
somite there arise three parts of
fundamental significance, viz., the sclerotome, the muscle plate,
and the cutis plate (dermatome), the primordium of the axial

Fig 107. — Transverse section through the twenty-ninth somite of a 29 s

embryo,
n. Cr., Neural crest. Neph., Nephrotome. W. D., Wolffian duct. Other
abbreviations as before.

skeleton, the voluntary muscles (excepting those of the head),

and derma respectively. The manner of origin of these parts
may be studied fully in an embryo of 25 to 30 somites, by com-
paring the most posterior somites, in which the process is begin-
ning, with somites of intermediate and anterior positions in the

series, which show successively later stages.

Figs. 107, 108, 109, and 110 represent transverse sections
through the twenty-ninth, twenty-sixth, tw^entieth, and seven-
teenth somites of a 29 s embryo. In the twenty-ninth somite
186 THE DEVELOPMENT OF THE CHICK

(Fig. 107) the primitive relations of the parts are still preserved.
In the twenty-sixth somite (Fig. 108) it will be seen that the
cells of the core and of the ventral and median wall of the somite
extending from the nephrotome to about the center of the neural
tube are becoming mesenchymal; they spread out towards the
notochord and the space between the latter and the dorsal aorta.
These cells constitute the sclerotome. The muscle plate extends
from the dorsal edge of the sclerotome to the dorso-median angle
of the wall of the somite, and the dermatome from this point
to the nephrotome.

nSr
Mj/.

'j^-
..ft ^^jst'-i=^-!^
^^^.:^A.
S26'.
~\
Sc/er

l^cp

-,£i-iv5'^

^j3;^v
Coe/.

'A/ep/i. ;«g9^_,
/;.//.

Fig. 108. — Transverse section through the twenty-sixth somite of a 29 s

embryo. (Same embryo as Fig. 107.)
Derm., Dermatome. My., Myotome. Scler., Sclerotome. V. c. p., Pos-
terior cardinal vein. Other abbreviations as before.

Fig. 109 is a section through the twentieth somite of the same

embryo. The sclerotome mesenchymal, and its cells
is entirely
are extending between the notochord and aorta, and along the
sides of the neural tube. The muscle-plate has now bent over
so that its inner surface is being applied against the dermatome,
but there is still a considerable cavity (myocoele) between the
two, at the lateral angle of the dermo-myotomic plate. The
lateral edge of the dermatome is freed from the nephrotome, and
turns in to a slight extent. Other details are readily understood
from the figure.
The growth of the free edge of the muscle-plate towards the
free lateral edge of the dermatome continues as illustrated in
FROM TWELVE TO THIRTY-SIX SOMITES 187

,^<V.,A.

"<

-iff -'-•^r o,<>«>-9^

-1^

"Si

.^s'sd^!^^

n
y h5*

: V
7
188 THE DEVELOPMENT OF THE CHICK

Figs. 109 and 110, until complete union of the two takes place
(Fig. Ill) and there is established a complete dermo-myotomic
plate in each somite. This is usually known as the myotome,
which therefore includes two layers; the external cutis-plate
or dermatome, and the internal muscle-plate. With the eleva-
tion of the axis of the body, the myotome gradually assumes
a nearly vertical position.

C/zor Jy^.

Fig. 110. — Transverse section through the seventeenth somite of a 29 s

embryo. (Same embryo as Fig. 107.)
am. Cav., Amniotic cavity. E. E. B. C, Extra-embryonic body-cavity.
Gn., Ganghon. mes'n. V., Mesonephric vesicle. S.-Am., Sero-amniotic con-
nection. Other abbreviations as before.

Other details concerning the early history of the sclerotome

are given in Chapter XIII, and it remains to add here only a short
description of certain changes in the cells of the myotome (myo-
blasts). In longitudinal sections the cells of the myotome are
seen to become spindle-shaped soon after the folding towards
the dermatome begins. The nuclei of the myoblasts are large
and stain less deeply than those of adjoining tissues. They
become elliptical in correspondence with the form of the cell-
bodies. Each myoblast soon stretches from anterior to pos-
terior faces of the somite, and this represents the first stage in
the differentiation of the voluntary muscles.
In later stages the myotomes send outgrowths into the limb-
buds and ventral body-wall for the formation of the voluntary
FRO:\r TWELVE TO THIRTY-SIX SOMITES 189