Text Book of Biology, Part 1: Vertebrata

by H. G. Wells

Section 24. We pointed out that the presence of the yolk in the frog's egg led to a difference in the size of the cells at the animal and vegetable poles. The late F.M. Balfour, borrowing a mathematical technicality, suggested that the rate of segmentation in any part of an ovum varies inversely with the amount of yolk. In the fowl's egg, except just at the germinal area, the active protoplasm is at a minimum, the inert yolk at a maximum; the ratio of yolk to protoplasm is practically infinity, and the yolk therefore does not segment at all. The yolk has diluted the active protoplasm so much as to render its influence inappreciable. The germinal area segments, and lies upon the yolk which has defeated the efforts of its small mingling of protoplasm to divide. Such a type of segmentation in which only part of the ovum segments is called meroblastic. If we compare this with the typical blastosphere of the lower type, we see that it is, as it were, flattened out on the yolk. This stage is shown in section in the lower figure of [Figure 1]. b.d., the blastoderm, is from this point of view, a part of the ripped and flattened blastosphere, spread out on the yolk; s.c. is the segmentation cavity, and y. the yolk.

Section 25. There is no open invagination of an archenteron in the fowl, as in the frog--, the gastrula, like the blastosphere, stage is also masked. But, in the hinder region of the germinal area, a thick mass of cells, grows inward and forward, and, appearing in the dorsal view of the egg as a white streak, is called the primitive streak (p.s.). By a comparison of the figures of frog and fowl the student will easily perceive the complete correspondence of the position of this with the blastopore of the frog. The relation of the two will be easily understood if we compare the fowl's archenteron to a glove-finger under pressure-- its cavity is obliterated-- and the frog's to the glove-finger blown out. The tension of the protoplasm, straining over the enormous yolk, answers to the pressure. The gastrula in the fowl is solid. The primitive streak is, in fact, the scar of a closed blastopore. As we should expect from this view of its homology, at the primitive streak, the three embryonic layers are continuous and indistinguishable ([Figure 2]). Elsewhere in the blastoderm they are distinctly separate. Just as the yolk cells of the frog form the ventral wall of the intestine, so nuclei appear along the upper side of the yolk of the fowl, where some protoplasm still exists, and give rise to the ventral hypoblastic cells. By conceiving a gradually increasing amount of yolk in the hypoblastic cells in the ventral side of the archenteron, the substantial identity of the gastrula stage in the three types, which at first appear so strikingly different, will be perceived. Carry [Figures 4 and 5] of the frog one step further by increasing the size of the shaded yolk and leaving it unsegmented, and instead of ar. in 5 show a solid mass of cells, and the condition of things in the fowl would at once be rendered.

Section 26. [Figure 3a] of the fowl will conveniently serve for comparison with [Figure 7] of the frog. The inturning of the medullary groove is entirely similar in the two cases. The mesoblast appears as solid mesoblastic somites. In the section above [Figure 4] this layer is shown as having split into somatopleur (so.) and splanchnopleur (spch.). [Figure 3] answers to [Figure 6] of the frog, and [Figure 4] is a later stage, in which the medullary groove is beginning to close at its middle part. The clear club-shaped area around the embryo (a.p.) is the area pellucida; the larger area without this is the area opaca (a.o.), in which the first bloodvessels arise by a running together and a specialization of cells. The entire germinal area grows steadily at its edges to creep over and enclose the yolk.

Section 27. So far, the essential differences between the development of fowl and frog, the meroblastic segmentation, absence of a typical gastrula, and the primitive streak, seem comprehensible on the theory that such differences are due to the presence of an enormous amount of yolk. Another difference that appears later is that, while the tadpole has an efficient pronephros, the fowl, which has no larval (free imperfect) stages in its life history, has the merest indication of such a structure.

Section 28. Another striking contrast, due to, or connected with, this plethora of yolk, is the differentiation of a yolk sac (= umbilical vesicle) and the development of two new structures, the amnion and allantois, in the fowl. If the student will compare [Figure 10] of the frog, he will see that the developing tadpole encloses in its abdomen all the yolk provided for it. This is a physical impossibility in the fowl. In the fowl (Figure 2, [Sheet 24]) the enormous yolk (Y.) lies outside of the embryo, and, as the cells of the germinal area grow slowly over it, umbilical bloodvessels are developed to absorb and carry the material to the embryo. In the case of an embryo sinking in upon, as it absorbs, this mass of nutritive material, a necessity for some respiratory structure is evident. From the hinder end of the fowl's intestine, in a position corresponding to the so-called, urinary bladder of the frog, a solid outgrowth, the allantois, which speedily becomes hollow, appears. Early stages are shown in Figures 1 and 2, [Sheet 24] (al.); while the same thing is shown more diagrammatically on [Sheet 23], Figure 6 (all.). This becomes at last a great hollow sac, which is applied closely to the porous shell, and the extent of which will be appreciated by looking at Figure 5, [Sheet 24], where the allantois is shaded. Allantoic bloodvessels ramify thickly over its walls, and aeration occurs through the permeable shell.

Section 29. The nature of the amnion will be understood by following Figures 4b, 5, and 6 on [Sheet 23]. The three embryonic layers are indicated by broken lines, dots, and black lines, just as they are in the frog diagrams. Not only is the embryo slowly pinched off from the yolk sac (y.s.), but, as the yolk is absorbed beneath it, and it grows in size, it sinks into the space thus made, the extra-embryonic somatopleur and epiblast rise up round it as two folds, which are seen closing in [5], and closed in [6], over the dorsal side of the young chick. In this way a cavity, a., lined by epiblast, and called the amniotic cavity, is formed. Dorsal to this, in [6,] comes a space lined by somatic mesoblast, and continuous with p.p., the pleuro-peritoneal cavity, or body cavity of the embryo. Outside this, again, is a layer, of somatopleur internally and epiblast externally, the false amnion (f.a.), which is continuous with the serous membrane (s.m.) enclosing the rest of the egg. The student should, carefully copy these diagrams, with coloured pencils or inks for the different layers, and should compare them with the more realistic renderings of Figures 2, 5, and 8, [Sheet 24].

Section 30. The heart in the fowl appears first as a pair of vessels, which unite to form a straight trunk in the median line, as the flattened-out embryo closes in from the yolk. The way in which this straight trunk is thrown, first of all, into the S shape of the fish heart, and then gradually assumes the adult form, is indicated roughly by [Figure 3]. In one respect the development of the heart does not follow the lines one would expect. Since, between the fish and the higher form comes the condition of such an animal as the frog, in which the auricles are divided, while there is only one ventricle, we might expect a stage in which the developing chick's heart would have one ventricle, and a septum between the auricles. But, as a matter of fact, the ventricles in fowl and rabbit are separated first, and the separation of the auricles follows, and is barely complete at birth.

Section 31. Two vitelline veins from the yolk sac (v.v.) flow into the heart from behind, as shown in [Figure 1]. A later more complete and more diagrammatic figure of the circulation is seen in [Figure 7]. At first there are two anterior cardinal (a.c.), and two posterior cardinal veins (p.c.) uniting to form Cuvierian sinuses (c.s.) that open into the heart just as in the dog-fish. But later the inferior cava is developed and extends backward, the posterior cardinals atrophy, the Cuvierian sinuses become the superior cavae, and the anterior cardinals the internal jugular veins. The vitelline veins (v.v.) flow, at first, uninterruptedly through the liver to the inferior cava, but, as development proceeds, a capillary system is established in the liver, and the through communication, the ductus venosus, is reduced-- at last-- completely. Bearing in mind that the yolk is outside the body in the fowl and inside it in the frog, the vitelline veins of the former have a considerable resemblance in position, and in their relation to the portal vein, to a portion of the single anterior abdominal vein. Blood is taken out to the allantois, however, by the arteries of the latter type.

Section 32. Five aortic arches are generally stated to appear altogether in the fowl, but not simultaneously. The first two, the mandibular and the hyoid vascular arches, early disappear, and are not comparable to any in the frog. The third is the first branchial arch, and, like the corresponding arch in the frog, forms the carotid artery; the second branchial is the aortic arch; and what has hitherto been regarded as the third (the fifth arch, i.e.) the pulmonary artery. A transitory arch, it is now known, however, appears between the second branchial and the last, and it is therefore the fourth branchial arch which is the pulmonary, just as it is in the frog.

Section 33. Blood, it may be mentioned, first appears in the area vasculosa, the outer portion of the area opaca. Embryonic cells send out processes, and so become multipolar; the processes of adjacent cells coalesce. The nucleus divides, and empty spaces appear in the substance of each of the cells.