The Works of Francis Balfour 1-3
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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.
Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
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Vol I. Separate Memoirs (1885)
III. On the Disappearance of the Primitive Groove in the Embryo Chick
With Plate I, figs. 68 and 1319.
THE investigations of Dursy (Der Primitivstreif des Hiihnchens, von Dr E. Dursy. Lahr, 1866) on the primitive groove, shewing that it is a temporary structure, and not connected with the development of the neural canal, have in this country either been ignored or rejected. They are, nevertheless, perfectly accurate ; and had Dursy made use of sections to support his statements I do not think they would so long have been denied. In Germany, it is true, Waldeyer has accepted them with a few modifications, but I have never seen them even alluded to in any English work. The observations which I have made corroborating Dr Dursy may, perhaps, under these circumstances be worth recording.
After about twelve hours of incubation the pellucid area of a hen's egg has become somewhat oval, with its longer axis at right angles to the long axis of the egg. Rather towards the hinder (narrower) end of this an opaque streak has appeared, with a somewhat lighter line in the centre. A section made at the time shews that the opaque streak is due partly to a thickening of the epiblast, but more especially to a large collection of the rounded mesoblast cells, which along this opaque line form a thick mass between the epiblast and the hypoblast. The mesoblast cells are in contact with both hypoblast and epiblast, and appear to be fused with the latter. The line of junction between them can, however, almost always be made out.
Soon after the formation of this primitive streak a groove is formed along its central line by a pushing inwards of the epiblast.
- 1 From the Qziarterly Journal of Microscopical Science, Vol. Kill, 1873.
The epiblast is not thinner where it lines the groove, but the mass of mesoblast below the groove is considerably thinner than at its two sides. This it is which produces the peculiar appearance of the primitive groove when the blastoderm is viewed by transmitted light as a transparent line in the middle of an opaque one.
This groove, as I said above, is placed at right angles to the long axis of the egg, and nearer the hind end, that is, the narrower end of the pellucid area. It was called " the primitive groove " by the early embryologists, and they supposed that the neural canal arose from the closure of its edges above. It is always easy to distinguish this groove, in transverse sections, by several well-marked characters. In the first place, the epiblast and mesoblast always appear more or less fused together underneath it ; in the second place, the epiblast does not become thinner where it lines the groove ; and in the third place, the mesoblast beneath it never shews any signs of being differentiated into any organ.
As Dursy has pointed out, there is frequently to be seen in fresh specimens, examined as transparent objects, a narrow opaque line running down the centre of this groove. I do not know what this line is caused by, as there does not appear to be any structural feature visible in sections to which it can correspond.
From the twelfth to the sixteenth hour the primitive groove grows rapidly, and by the sixteenth hour is both absolutely and considerably longer than it was at the twelfth hour, and also proportionately longer as compared with the length of the pellucid area.
There is a greater interval between its end and that of the pellucid area in front than behind.
At about the sixteenth hour, or a little later, a thickening of the mesoblast takes place in front of the primitive groove, forming an opaque streak, which in fresh specimens looks like a continuation from the Anterior extremity of the primitive groove (vide PI. I, fig. 8). From hardened specimens, however, it is easy to see that the connection of this streak with the primitive groove is only an apparent one. Again, it is generally possible to see that in the central line of this streak there is a narrow groove. I do not feel certain that there is no period when this groove may not be present, but its very early appearance has not been recognized either by Dursy or by Waldeyer. Moreover, both these authors, as also His, seem to have mistaken the opaque streak spoken of above for the notochord. This, however, is not the case, and the notochord does not make its appearance till somewhat later. The mistake is of very minor importance, and probably arose in Dursy's case from his not sufficiently making use of sections. At about the time the streak in front of the primitive groove makes its appearance a semicircular fold begins to be formed near the anterior extremity of the pellucid area, against which the opaque streak, or as it had, perhaps, better be called, " the medullary streak," ends abruptly.
This fold is the head fold, and the groove along the medullary streak is the medullary groove, which subsequently forms the cavity of the medullary or neural canal.
Everything which I have described above can without difficulty be made out from the examination of fresh and hardened specimens under the simple microscope ; but sections bring out still more clearly these points, and also shew other features which could not have been brought to light without their aid. In PL I, figs. 6 and 7, two sections of an embryo of about eighteen hours are shewn. The first of these passes through the medullary groove, and the second of them through the extreme anterior end of the primitive groove. The points of difference in the two sections are very obvious.
From fig. 6 it is clear that a groove has already been formed in the medullary streak, a fact which was not obvious in the fresh specimen. In the second place the mesoblast is thickened both under the groove and also more especially in the medullary folds at the sides of the groove ; but shews hardly a sign of the differentiation of the notochord. So that it is clear that the medullary streak is not the notochord, as was thought to be the case by the authors above mentioned. In the third place there is no adhesion between the epiblast and the mesoblast. In all the sections I have cut through the medullary groove I have found this feature to be constant; while (for instance, as in PL I, figs. 7, 9, 17) all sections through the primitive groove shew most clearly an adhesion between the epiblast and mesoblast. This fact is both strongly confirmatory of the separate origins of the medullary and primitive grooves, and is also important in itself, as leaving no loophole for supposing that in the region of embryo there is any separation of the cells from the epiblast to form the mesoblast.
By this time the primitive groove has attained its maximum growth, and from this time begins both absolutely to become smaller, and also gradually to be pushed more and more backwards by the growth of the medullary groove.
The specimen figured in PI. I, fig. 18, magnified about ten diameters, shews the appearance presented by an embryo of twenty-three hours. The medullary groove (me) has -become much wider and deeper than it was in the earlier stage ; the medullary folds (A) are also broader and more conspicuous. The medullary groove widens very much posteriorly, and also the medullary folds separate far apart to enclose the anterior end of the primitive groove (pr\
All this can easily be seen with a simple microscope, but the sections taken from the specimen figured also fully bear out the interpretations given above, and at the same time shew that the notochord has at this age begun to appear. The sections marked 13 17 pass respectively through the lines with corresponding numbers in fig. 18. Section I (fig. 13) passes through the middle of the medullary canal.
In it the following points are to be noted, (i) That the epiblast becomes very much thinner where it lines the medullary canal (me), a feature never found in the epiblast lining the primitive groove. (2) That the mesoblast is very much thickened to form the medullary folds at A, A, while there is no adherence between it and the epiblast, below the primitive groove. (3) The notochord (c/i) has begun to be formed, though its separation from the rest of the mesoblast is not as yet very distinct 1 .
In fig. 14 the medullary groove has become wider and the medullary folds broader, the notochord has also become more expanded : the other features are the same as in section I. "In the third section (fig. 15) the notochord is still more expanded; the bottom of the now much expanded medullary groove has become raised to form the ridge which separates the medullary from the primitive groove. The medullary folds are also flatter and broader than in the previous section. Section 4 (fig. 16) passes through the anterior end of the primitive groove. Here the notochord is no longer visible, and the adherence between the mesoblast and epiblast below the primitive groove comes out in marked contrast with the entire separation of the two layers in the previous sections.
- 1 In the figure the notochord has been made too distinct.
The medullary folds (A) are still visible outside the raised edges of the primitive groove, and are as distinctly as possible separate and independent formations, having no connection with the folds of the primitive groove. In the last section (fig. 17), which is taken some way behind section 4, no trace of the medullary folds is any longer to be seen, and the primitive groove has become deeper. This series of sections, taken in conjunction with the specimen figured in fig. 1 8, must remove all possible doubt as to the total and entire independence of the primitive and medullary grooves. They arise in different parts of the blastoderm ; the one reaches its maximum growth before the other has commenced to be formed ; and finally, they are distinguished by almost every possible feature by which two such grooves could be distinguished.
Soon after the formation of the notochord, the proto-vertebrse begin to be formed along the sides of the medullary groove (PI. I, fig. 19, pv). Each new proto-vertebra (of those which are formed from before backwards) arises just in front of the anterior end of the primitive groove. As growth continues, the primitive groove becomes pushed further a"nd further back, and becomes less and less conspicuous, till at about thirty-six hours only a very small and curved remnant is to be seen behind the sinus rhomboidalis ; but even up to the forty-ninth Dursy has been able to distinguish it at the hinder end of the embryo.
The primitive groove in the chick is, then, a structure which appears very early, and soon disappears without entering directly into the formation of any part of the future animal, and without, so far as I can see, any function whatever. It is clear, therefore, that the primitive groove must be the rudiment of some ancestral feature ; but whether it is a rudiment of some structure which is to be found in reptiles, or whether of some earlier form, I am unable to decide. It is just possible that it is the last trace of that involution of the epiblast by which the hypoblast is formed in most of the lower animals. The fact that it is formed in the hinder part of the pellucid area perhaps tells slightly in favour of this hypothesis, since the point of involution of the epiblast not unfrequently corresponds with the position of the anus.
EXPLANATION OF PLATE I. Figs. 68 and 1319.
Figs. 6 and 7 are sections through an embryo rather earlier than the one drawn in fig. 8. Fig. 6 passes through the just commencing medullary groove (md), which appears in fresh specimens, as in fig. 8, merely as an opaque streak coming from the end of the primitive groove. The notochord is hardly differentiated, but the complete separation of mesoblast and hypoblast under the primitive groove is clearly shewn. Fig. 7 passes through the anterior end of the primitive groove (pr), and shews the fusion between the mesoblast and epiblast, which is always to be found under the primitive groove.
Fig. 8 is a view from above of a twenty hours' blastoderm, seen as a transparent object. Primitive groove (pr). Medullary groove (md}, which passes off from the anterior end of the primitive groove, and is produced by the thickening of the mesoblast. Headfold (//).
Figs. 13 17 are sections through the blastoderm, drawn in fig. 18 through the lines i, 2, 3, 4, 5 respectively.
The first section (fig. 13) passes through the true medullary groove (me); the two medullary folds (A, A) are seen on each side with the thickened mesoblast, and the mesoblast cells are beginning to form the notochord (nc) under the medullary groove. There is no adherence between the mesoblast cells and the epiblast under the medullary groove.
The second (fig. 14) section passes through the medullary groove where it has become wider. Medullary folds, A, A ; notochord, ch.
In the third section (fig. 15) the notochord (ch) is broader, and the epiblast is raised in the centre, while the medullary folds are seen far apart at A.
In section fig. 16 the medullary folds (A) are still to be seen enclosing the anterior end of the primitive groove (pr). Where the primitive groove appears there is a fusion of the epiblast and mesoblast, and no appearance of the notochord.
In the last section, fig. 1 7, no trace is to be seen of the medullary folds.
Figs. 18 and 19 are magnified views of two hardened blastoderms. Fig. 18 is twenty-three hours old; fig. 19 twenty-five hours. They both shew how the medullary canal arises entirely independently of the primitive groove and in front of it, and also how the primitive groove gets pushed backwards by the growth of the medullary groove, pv, Proto-vertebrae ; other references as above. Fig. 1 8 is the blastoderm from which sections figs. 13 17 were cut.
Cite this page: Hill, M.A. (2021, October 22) Embryology The Works of Francis Balfour 1-3. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/The_Works_of_Francis_Balfour_1-3
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