The Origin of Vertebrates

by Walter Holbrook Gaskell

From this description it is clear that the vascular supply of the branchial segment of Ammocœtes would resemble most closely the vascular supply of the Limulus branchial appendage, if the ventral aorta of the former was derived from two longitudinal veins, homologous with the paired longitudinal venous sinuses of the latter.

A priori, such a derivation seems highly improbable; and yet it is precisely the manner in which embryology teaches us that the heart and ventral aorta of the vertebrate have arisen.

The Origin of the Invertebrate Heart and the Origin of the Vertebrate Heart.

Not only does the vertebrate heart differ from that of the invertebrate, in that it is branchial while the latter is systemic, but also it is unique in its mode of formation in the embryo. In the Appendiculata the heart is formed as a single organ in the mid-dorsal line by the growth of the two lateral plates of mesoblast dorsalwards, the heart being formed where they meet. In Mammalia and Aves, the heart and ventral aorta commence as a pair of longitudinal veins, one on each side of the commencing notochord.

If the embryo be removed from the yolk, the surface of the embryo covering these two venous trunks can be spoken of as the ventral surface of the embryo at that stage, and indeed we find that in the present day there is an increasing tendency to speak of this surface as the ventral surface of the embryo. Thus, Mitsukuri, in his studies of chelonian embryos, lays great stress on the importance of surface views and when the embryo has been removed from the yolk, figures and speaks of its ventral surface. So, also, Locy and Neal find that the best method of seeing the early segments of the embryo is to remove the embryo from the yolk, and examine what they speak of as a ventral view. At the period, then, before the formation of the throat, we may say that on the ventral surface of the embryo a pair of longitudinal venous sinuses are found, one on each side of the mid-ventral line, which are in the same position with respect to the mid-axis of the embryo as are the longitudinal venous sinuses in Limulus.

The next step is the formation of the throat by the extension of the layers of the embryo laterally to meet in the mid-line and so form the pharynx, with the consequence that a new ventral surface is formed; these two veins, as is well known, travel round also, and, meeting together in the new mid-ventral line, form the subintestinal vein, the heart, and the ventral aorta.

What is true of Mammalia and Aves, has been shown by P. Mayer to be true universally among vertebrates, so that in all cases the heart and ventral aorta have arisen by the coalescence in the new mid-ventral line of two longitudinal venous channels, which were originally situated one on each side of the notochord, in what was then the ventral surface of this part of the embryo. This history is especially instructive in showing how the pharyngeal region is formed by the growing round of the lateral mesoblast, i.e. the muscular and other mesoblastic tissues of the branchial segments, and how the two longitudinal veins take part in this process. The phylogenetic interpretation of this embryological fact seems to be, that the new ventral surface of the vertebrate in this region is formed, not only by the branchial appendages, but also by the growth ventrally of that part of the original ventral surface which covered each longitudinal venous sinus.

The following out of the consecutive clues, which one after the other arise in harmonious succession as the necessary sequence of the original working hypothesis, brings even now into view the manner in which the respiratory portion of the alimentary canal arose, and gives strong hints as to the position of that part of the arthropod which gave origin to the notochord. Here I will say no more at present, for the origin of the new alimentary canal of the vertebrate and of the notochord will be more fittingly discussed as a whole, after all the other organs of the vertebrate have been compared with the corresponding organs of the arthropod.

Fig. 72.—Diagram (Upper Half of Figure) of the Original Position of Veins (H) which come together to form the Heart of a Vertebrate.