Paper - A Young Human Embryo (Embryo Dobbin) with Head-Process and Prochordal Plate
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Hill, J. P. and Florian, J., 1931. A Young Human Embryo (Embryo Dobbin) with Head-Process and Prochordal Plate. Phil. Tran. Roy. Soc. London B, 219, 443-486.
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By J. P. HILL, D.Sc., F.R.S., and J. FLORIAN, M .D.
(From the Department of Anatomy and Embryology, University College, London.)
(Received July 27, 1931.- Read November 19, 1931.)
Our knowledge of the early development of Man has made marked progress during recent years, numerous fairly Well—preserved embryos having been described in greater or less detail. There are still, however, numerous gaps in our knowledge, pertaining not merely to points of detail but to fundamental questions, so that any early embryo reasonably Well preserved and reasonably Well sectioned is deserving of careful study.
Our specimen belongs to the comparatively late presomite stage represented in the literature by such embryos as K113 (GROSSER, 1913), Wa17 (GROSSER, 1931), Peh.1- Hochstetter (ROSSENBECK, 1923), and the embryo of INGALLS (1918), and characterised by the presence of an elongated primitive streak, a luminated chorda-process and a prochordal plate. We offer no apology, however, for presenting a fairly detailed, description of yet another example of this stage, since our embryo amongst other things provides a more complete picture of the cranial region of the head~process than any specimen hitherto described.
History and Treatment of the Specimen
The chorionic vesicle, the embryo of which forms the subject-matter of this paper, was presented to one of us (HILL) by Dr. ROY DOBBIN, of Cairo, through the kind ofﬁces of Professor D. E. DERBY. In appreciation of his valuable gift, We have much pleasure in associating Dr. DOBBIN’s name with the embryo.
The clinical history supplied by Dr. DOBBIN is as follows: “ Coitus, 6.10.23 ; effort probably causing abortion, 21.10.23; ﬁrst bleeding, 22.10.23; abortion (painless), 23.10.23."
Although an abortion, we see no reason to regard the specimen as other than perfectly normal. The chorionic vesicle (which was preserved in spirit) was, when received, somewhat ﬂattened and shrunken (ﬁg. 1, Plate 29). Except over a small area on one side (approximately 3 X 2 mm. in diameter), which was almost bare, the vesicle possessed a fairly uniform covering of short, close~set, branched villi (ﬁg. 2, Plate 29), to which at one point a small fragment of blood—clot adhered. Including the villi, its dimensions in alcohol were as follows : 11 - 5 mm. (in long diameter) X 8 -5 mm. (in short diameter) X 45 mm. (in thickness). After clearing in oil of cedar-wood, the corresponding internal diameters were 9 mm. X 5-5 mm. X 2-5 mm. 2 The vesicle, after being photographed and drawn, was dehydrated and cleared in oil of cedar-wood. A small portion of the chorion, including the bare area, was then carefully removed, and through the opening so made it was possible, fortunately enough, to locate the embryo under the binocular dissecting microscope. The embryo was then isolated along with the segment of the chorion to which it was attached, and stereo-photographs were successfully taken of it, in the cleared condition in oil of cedar—wood. Subsequently Mr. A. K. MAXWELL, with the aid of these photographs and the camera lucida, made the beautiful drawings representing the left lateral and dorsal aspects of the embryo which are reproduced as ﬁgs. 3 and 4, Plate 29. These ﬁgures are, we believe, unique, in that they are the only illustrations extant of the dorsal andlateral views of the human embryo at this particular phase of development which have been made directly from the embryo itself, and not from models or reconstructions.
The embryo and the related piece of chorion were double—embedded in cedar-wood oil-pyroxelene and paraffin, and cut cranio—caudally by H. BARKER into a complete and really very ﬁne series of sections at 8 (1., the sectional plane being almost exactly transverse to the long axis of the embryo. The state of preservation of the latter proved to be by no means perfect cytologically, but is sufficiently good to ustify us in giving a fairly detailed account of its structural condition. The most obvious ﬁxation—defect is the partial disintegration of the most cranial portion of the Shield-ectoderm, the granular detritus resulting therefrom lying partly in the amniotic cavity and partly below the ectoderm, in a space enclosed between the latter and the detached basement membrane. The fact that the cranial region of the early embryo is the ﬁrst part to undergo dissolution seems to be Well recognised. Inspection of ﬁg. 3, Plate 29, will show that the embryo no longer occupies its normal position in relation to the chorion, but has been displaced in the ventral direction. Fortunately the deformation accom- panying this displacement has affected only the most caudal part of the embryo and the connecting stalk, and is not of a serious character.
The following measurements were made whilst the embryo was still in oil of cedar- Wood, but must be regarded as approximate only. The lettering refers to text—ﬁg. 2 :-
|Anterior margin, embryonal shield to region of cloacal membrane (A-Cl.)||0.98 mm|
|Vertical diameter (D-F) of yolk-sac||1.092 mm|
|Antero-posterior diameter of yolk-sac, near its mid-region||0.98 mm|
|Vertical height, amnio-embryonal vesicle (D-E)||0.468 mm|
|Length of yolk-sac process||0.88 mm|
The measurements of the embryo based on the sections and graphic reconstructions are set forth, along with those of other early embryos, in the tables provided at the end of this paper (p. 480-81).
We take this opportunity of expressing our very grateful thanks to Dr. ROY DOBBIN and to Professor D. E. DERBY for the gift of this interesting embryo. We are further greatly indebted to Professor J. S. B. STOPFORD for the loan of the presomite Manchester embryo, No. 1285 ; to Professor J. C. BRASH for the opportunity of examining the sections of the Thompson-Brash embryo; to Hofrat Professor F. HOCHSTETTER for permission to study the Peh.,-Hochstetter embryo, described by ROSSENBECK, 1923 ; and to Dr. O. BITTMANN, Brno, for the gift to one of us (FLORIAN) of the Bi(ttmann) 24 embryo, to which reference is made on pp. 466—7 . We desire also to thank Mr. A. K. MAXWELL, Artist to the Department, for the care and skill he has expended on the illustrations on Plates 29-34, Which, with the exception of ﬁgs. 1 and 2, are all based on photomicrographs; and to Mr. F. J. PITTOCK for his invaluable assistance in photography.
The Embryo as a Whole
The embryonal body, represented by the two vesicles——the amnio-embryonal and yolk—sac vesicles-——-is attached by a very distinct connecting stalk to the chorion (ﬁg. 3, Plate 29). It will be observed that, as has just been mentioned, the dorsal surface of the amnio~embryonal vesicle does not face the chorion, as is the rule in this stage of development, the embryonal shield forming an angle of about 125° with the inner surface of that membrane. This is evidently due to the artiﬁcial displacement of the embryonic body, including the connecting stalk, in the ventral direction. At the same time the embryo and the connecting stalk have suﬁered a slight rotation round the long axis of the body towards the left side (as may be seen from fig. 4, Plate 29). As is usual at this stage, the yolk—sac vesicle is distinctly larger than the amnio-embryonal.
Text-Fig. 1. Graphic reconstruction of the dorsal view of the embryonic formation; Dobbin embryo X 75. - Outlines of the embryonal shield (E.s.), yolk-sac (Y.s.) and connecting stalk (c.st.) indicated by a thick line-; outlines of the allantoic canal (ALI = tubular part, AL2 =2: vesicular part) indicated by interrupted lines; outline of the caudal end of the amniotic cavity (Amn.) dotted; cloacal membrane (UL) lined longitudinally where indistinct, cross—hatched where distinct ; P7‘..S‘. =2 primitive streak; chorda process and chorda canal lined transversely; lateral mesodermal bands lined transversely and dotted; ventral openings of the chorda canal marked by arrows on the right, its dorsal opening by an arrow on the left. ++ = HENSEN’s knot ; prochordal plate marked by large dots. On the scale is indicated the level of every ﬁfth section. The median plane is indicated by an arrow.
Text-Fig. 2. Graphic reconstruction of the median section through the Dobbin embryo X 7 5. - Ectoderm black; endoderm lined horizontally; primitive streak lined vertically; +=HENSEN’s knot; chorda-process lined obliquely; prochordal plate (Pr.pl.) lined horizontally and dotted; mesoderm dotted; cloacal membrane (O'l.) cross—hatched. Limits of the embryonal shield marked by arrows. AL; 2: proximal (tubular) part of the allantoic canal; Al.2 == its distal (vesicular) part. Hor. —-—=. horizontal plane (the plane of the projection in text-ﬁgs. 1 and 8). The caudal part of the embryonic formation, including the cloacal membrane, the allantoic canal, the caudal part of the d amniotic cavity and the connecting stalk are projected into the median plane, and their outlines are marked by interrupted lines. , The cell-layer underneath the intermediate region and the most cranial portion of the head-process, the signiﬁcance of Which is doubtful, is left White.
The embryonal shield forms in the lateral view (text-ﬁg. 2) an open S-shaped curve. It reaches cranially up to the contour-line formed by the amniotic and yolk—sac vesicles, and ends caudally just in front of the connecting stalk.
The sections show that the embryonal shield over its cranial two-thirds is not ﬂat as is normal at this stage, but is dorsally concave so that in the lateral view of the embryo (ﬁg. 3, Plate 29) the ventral surface of the left half of the embryonal shield is turned towards the observer and appears much darker than the right half, which is only partly visible.
The yolk—sac is roughly conical in form; its broad base underlies the embryonal shield, whilst its apical region narrows to form a spike-like projection. This latter is prolonged into a ﬁlamentous process (about 0 - 88 mm. in length) which exhibits three enlargements, an elongated thickening close to the apex of the yolk-sac, an oval swelling near the middle of its length and a quite minute nodule close to its termination (ﬁg. 3, Plate 29). It ended freely, without reaching the chorion. The entire process represents the yolk~sac process first described by STRAHL and BENEKE 1910, and subsequently observed in all adequately preserved early human embryos in a more or less developed condition. The process attains its maximum development in the embryos Teacher-Bryce II (BRYCE, 1924) and BENEKE, in which it reaches and is attached to the opposite wall of the chorion. The latest stage in which it has been observed in this complete condition is the embryo, Ct POLITZER, with seven pairs of somites (G.PoLI'rzER, 1930).
The yolk-sac at its postero-dorsal extremity is in the sections seen to be continued into the mesoderm of the connecting stalk in the form of an elongated canal (the allantoic canal). At its caudal end this enlarges to form a distinct thin-walled vesicle, which can be seen. both in the dorsal and lateral views of the embryo (ﬁgs. 3 and 4, Plate 29) as a small oval structure on the surface of the connecting stalk close to its attachment to the mesoderm of the chorion, and on the left side of the same 9 (cf. also ﬁgs. 28 and 29, Plate 32). p , P "
On the surface of the yolk-sac (ﬁg. 3, Plate 29) a number of vessel—primordia can be clearly observed.
In the dorsal view (text-ﬁg. 1 and ﬁg. 4, Plate 29), the shoe—sole shaped form of the embryonal shield is apparent. Along its middle line the head-process and the primitive groove could be made out in the intact specimen. The pointed prolongation of the amniotic cavity along the dorsal surface of the connecting stalk is very distinctly seen in the dorsal view, whilst the amniotic membrane itself, forming the roof of the amnio- embryonal vesicle, is seen in the lateral View (ﬁg. 3, Plate 29).
(i) Head-process and Prockordal Plate;-——In our preliminary note (HILL-FLORIAN, 1931) we distinguished three portions in the axial preblastoporic structures in our specimen. The most cranial portion we interpreted as a prochordal plate which had already produced a thick mass of prochordal mesoderm. Since the publication of our preliminary note one of us (F.), by the kind permission of Professor HOCHSTETTER, has been able to study the region in question in the embryo Peh.1-Hochstetter (RossEN~BECK, 1923). As this embryo is much better preserved in the region of the prochordal plate than the Dobbin embryo, we have been able to convince ourselves that what we have interpreted as prochordal plate in our preliminary note really represents the cranial end of the head~process. The actual prochordal plate (text-ﬁg. 8) in the Dobbin embryo is very indistinct because of the poor preservation of this region, and it is also possible that the plate varies in diﬁerent embryos. It is represented by an area of thickened endoderm in front of the cranial end of the head—process, presenting no distinct evidence of mesoderm formation. The thickened endoderm of this region can be traced continuously backwards underneath and laterally to the cranial part of the head-process up to section 27. Caudally it becomes indistinct. _ In the head—process itself we can distinguish three parts : (1) A caudal part forming about the caudal half of the entire formation and extending through sections 7 2-46; (2) a shorter part which extends forwards from the caudal segment and gradually widens to pass into continuity with the third part. This second portion we distin- guished in our preliminary note as the intermediate region. It is comprised in sections 46 to about 32; (3) a short but broad terminal or cranial part occupying sections 31 to 23.
The caudal part (1) is formed by a typical chorda—canal, its lumen communicating by seven ventral openings with the yolk—sac cavity and by a dorsal opening on the surface of I-IENsEN’s knot with the amnioticcavity. 'I‘he_intermediate part (2) is formed by a segment of the head—process, which here has undergone differentiation into a median chorda~process and lateral mesodermal bands. The most cranial part (3) (which we distinguished as prochordal plate in our preliminary note) is formed by a distinct, dense cellular mass, Olevoid of any differentiation.
The ectoderm of the amnio-embryonal vesicle is first encountered in section 18 and its cavity in section 19 (ﬁg. 9, Plate 30). Between the cranial end of the amnio-embryonal vesicle and the yolk—sac endoderm, there is present a sheet of mesoderm which can be traced cranially up to section 15. . Its cells are fairly compactly arranged and are connected by ﬁne cytodesmata with the ectoderm and endoderm. The median part of this mesodermal sheet passes into continuity behind ‘with the cranial tip of the head—process, which is first visible in section 23. In front of the head-process the prochordal plate appears. The first indication of a thickened endodermal area in the median region seems to occur in section 20, but is only slight. It broadens out in section 21 (fig. 30, Plate 32), and in section 22 becomes more distinct. In section 23 (fig. 31, Plate 32) the prochordal endoderm has almost the same structure as in section 22, the nuclei being in places several deep ; in this section the cranial tip of the head-process appears in the median plane, intercalated into the mesodermal sheet. The head-process rapidly increases in size as it is traced back, and already in section 25 has assumed the appearance seen in section 27 (fig. 10, Plate 30 ; fig. 32, Plate 32). These ﬁgures show clearly the mesoderm-like character of this part of the head-process. Laterally on each side, the endoderm is still ‘distinctly thickened, but below the head- process itself it is no longer distinguishable as a separate layer. The cells of the head- process are for the most part compactly arranged and form a mass of considerable thickness, which laterally passes over without deﬁnite limit into the adjoining thin sheets of primitive streak mesoderm. The cells contain deeply staining chromatophilic granules which resemble those described by BONNET (1901, p. 287) in the “ Erganzungs- platte ” of the dog, and they are also present in the minute intercellular spaces. In places the cells tend to be arranged round small cavities (one in section 24, two in section 27, ﬁg. 32,Plate 32, and one in section 28, ﬁg. 33, Plate 32), but these vesicles are too indistinct and too irregularly distributed to have any important signiﬁcance.
The thickened endoderm continued back from the prochordal plate can be traced on either side of the head-process up to section 27 and, as in the preceding sections, is fused with the head-process in the median line. Caudally to this section the thickened endoderm area becomes indistinct, and behind section 34 is certainly no longer present. The possible explanation. of these facts we shall discuss later in connection with the description of the same region in other human embryos (t. p. 482).
In section 30 (ﬁg. 34, Plate 32) the head-process has become reduced, both in thickness and in width, as compared with that of the preceding sections, whilst its cells present a somewhat looser appearance, and the chromatophilic granules have almost completely disappeared. The next section (N o. 31, ﬁg. 35, Plate 32) presents a very similar appearance, except that there are no granules. and the axial mass is more massive.
In both sections no distinct endoderm can be observed below the head-process.
Section 32 (ﬁg. 36, Plate 32) marks the beginning of our intermediate region, since in it we encounter the tip of the chorda-process in the form of a compact circular mass, situated asymmetrically on the left side and composed of cells radially arranged round a potential lumen and possessing peripheral nuclei. On the right of the chorda-process the mesoderm of the head-process forms a dense mass, but on the left it is quite diffuse. There is no clearly deﬁned underlying layer of endoderm.
In the following section (No. 33, ﬁg. 37, Plate 33) the chorda-process is represented by a distinct chorda—canal, with a very obvious lumen and with a mesodermal mass closely adjoining it on each side ; that on the left being more diﬁuse, that on the right more compact and much less massive than the mass in the corresponding position in the preceding section. Both the masses as well as the chorda-canal closely adjoin the endoderm, which is not distinct as a separate layer. We regard these lateral masses as part of the lateral mesodermal bands, characteristic of the intermediate region.
Similar conditions are observed in section 34 (ﬁg. 38, Plate 33), except that the lumen of the chorda—-canal is much reduced, and in section 35 (ﬁg. 39, Plate 33) only a trace of it remains. The endoderm below the axial region is not distinguishable as a separate layer. This is the case also in section 36 (ﬁg. 40, Plate 33), where the lumen of the chorda—canal again becomes quite distinct. I
In section 37 (fig. 41, Plate 33) the chorda-process is solid, and the limits of its ventral wall are difficult to determine. No endoderm is distinguishable below the head—process. The mesodermal bands are distinct, the right being more compact than the left, and are delimited from the chorda—-process dorsally by a groove. Section 38 (ﬁg. 42, Plate 33) is very similar, except that the right mesodermal band is distinctly larger than the left. It is evident that no endoderm is distinct below the head-process. A broad shallow groove (already indicated in the two preceding sections) is now to be found underneath the latter, in contrast to the ventral convexity in the corresponding position in the most cranial region of the head-process. It can be traced back into continuity with the well-marked median groove underlying the caudal half of the head»-process. e
In sections 39--43 (ﬁgs. 43--46, Plate 33) there is a continuous lumen in the chorda- canal; no deﬁnite endoderm is present below the median part of the head-process. The lateral mesodermal bands are symmetrically disposed and quite well marked.
In section 44 (ﬁg. 47, Plate 33) the chorda-canal, broader than in ﬁg. 46, does not show the typical characters seen in the preceding sections. Its lumen appears as a large eccentric cavity of somewhat irregular outline, whilst its cells exhibit an epithelial arrangement only over its dorsal half, those of its ventral wall being irregularly arranged and fused with the endoderm. The lateral mesodermal. bands are now much reduced in size. Essentially the same features are seen in the succeeding section (No. 45, ﬁg. 48, Plate 33). The cells exhibit an epithelial character only on the dorsal side of the solid chorda-process. The right mesodermal band is still distinct, but small, and the left has practically disappeared. In section 46 (ﬁg. 49, Plate 33) the sectional appearance of the axial formation has considerably altered. The chorda-process now appears more massive, owing to the compact groupingof its cells. They show practically no distinct epithelial arrangement, and are fused ventrally with the endoderm, which below the chorda-process is not distinguishable as an independent layer. There may possibly" still be a remnant of the right mesodermal band, but the left has completely disappeared. We may regard this section as marking the junction between our intermediate region and the caudal part of the axial formation, which, as we shall see, consists only‘ of a chorda—canal, devoid of lateral mesodermal bands. In section 47 (ﬁg. 50, Plate 33) the chorda-process is massive and takes the form of a chorda-canal semicircular in outline, the lumen of which appears as a very narrow ﬁssure with a wide opening below into the yolk—sac cavity, and whose wall is composed of cells exhibiting an epithelial arrangement. Marginally, on each side, the wall passes into continuity on the one hand with the lateral (primitive streak) mesoderm, and on the
- We employ the old term “ head—process ” for the axial formation which extends forwards from I-IENsEN’s knot, the homologue of the dorsal lip of the blastopore (notopore). This formation comprises not only the primordium of a part of the mesoderm (which does not seem to be very extensive), but also that of the chorda, and therefore such terms as “ Mesodermsackchen ” (HERTWIG, 0., 1906) or “ kranialer Mesoblastfortsatz ” (WALDEYER, 1929) are not suitable.
- Right and left in the description of the ﬁgures correspond to the reader’s right and left, but‘ actually to the left and right sides of the embryo respectively, 6.6., the apparent right side of the ﬁgure is the actual left in the embryo and vice versa. 3
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Published December 9, 1931. In Australia copyright has expired - creator died before 1955, provided; work was also published before 1955.
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