When we examine the embryology of the human nervous system, we must start from the important fact, which we have already seen, that the first structure of it in man and all the higher Vertebrates is the simple medullary tube, and that this separates from the outer germinal layer in the middle line of the sole-shaped embryonic shield. As the reader will remember, the straight medullary furrow first appears in the middle of the sandal-shaped embryonic shield. At each side of it the parallel borders curve over in the form of dorsal or medullary swellings. These bend together with their free borders, and thus form the closed medullary tube (Figs. 133–137). At first this tube lies directly underneath the horny plate; but it afterwards travels inwards, the upper edges of the provertebral plates growing together between the horny plate and the tube, joining above the latter, and forming a completely closed canal. As Gegenbaur very properly observes, “this gradual imbedding in the inner part of the body is a process acquired with the progressive differentiation and the higher potentiality that this secures; by this process the organ of greater value to the organism is buried within the frame.” (Cf. Figs. 143–146).
In the Cyclostoma—a stage above the Acrania—the fore end of the cylindrical medullary tube begins early to expand into a pear-shaped vesicle; this is the first outline of an independent brain. In this way the central marrow of the Vertebrates divides clearly into its two chief sections, brain and spinal cord. The simple vesicular form of the brain, which persists for some time in the Cyclostoma, is found also at first in all the higher Vertebrates (Fig. 153 hb). But in these it soon passes away, the one vesicle being divided into several successive parts by transverse constrictions. There are first two of these constrictions, dividing the brain into three consecutive vesicles (fore brain, middle brain, and hind brain, Fig. 154 v, m, h). Then the first and third are sub-divided by fresh constrictions, and thus we get five successive sections (Fig. 155).
Fig. 294–296—Central marrow of the human embryo from the seventh week, 4/5 inch long. (From Kölliker.) Fig. 294. The brain from above, v fore brain, z intermediate brain, m middle brain, h hind brain, n after brain. Fig. 2955. The brain with the uppermost part of the cord, from the left. Fig. 296. Back view of the whole embryo: brain and spinal cord exposed.
In all the Craniotes, from the Cyclostoma up to man, the same parts develop from these five original cerebral vesicles, though in very different ways. The first vesicle, the fore brain (Fig. 155 v), forms by far the largest part of the cerebrum—namely, the large hemispheres, the olfactory lobes, the corpora striata, the callosum, and the fornix. From the second vesicle, the intermediate brain (z), originate especially the optic thalami, the other parts that surround the third cerebral ventricle, and the infundibulum and pineal gland. The third vesicle, the middle brain (m), produces the corpora quadrigemina and the aqueduct of Sylvius. From the fourth vesicle, the hind brain (h), develops the greater part of the cerebellum—namely, the vermis and the two small hemispheres. Finally, the fifth vesicle, the after brain (n), forms the medulla oblongata, with the quadrangular pit (the floor of the fourth ventricle), the pyramids, olivary bodies, etc.
We must certainly regard it as a comparative-anatomical and ontogenetic fact of the greatest significance that in all the Craniotes, from the lowest Cyclostomes and fishes up to the apes and man, the brain develops in just the same way in the embryo. The first rudiment of it is always a simple vesicular enlargement of the fore end of the medullary tube. In every case, first three, then five, vesicles develop from this bulb, and the permanent brain with all its complex anatomic structures, of so great a variety in the various classes of Vertebrates, is formed from the five primitive vesicles. When we compare the mature brain of a fish, an amphibian, a reptile, a bird, and a mammal, it seems incredible that we can trace the various parts of these organs, that differ so much internally and externally, to common types. Yet all these different Craniote brains have started with the same rudimentary structure. To convince ourselves of this we have only to compare the corresponding stages of development of the embryos of these different animals.
This comparison is extremely instructive. If we extend it through the whole series of the Craniotes, we soon discover this interesting fact: In the Cyclostomes (the Myxinoida and Petromyzonta), which we have recognised as the lowest and earliest Craniotes, the whole brain remains throughout life at a very low stage, which is very brief and passing in the embryos of the higher Craniotes; they retain the five original sections of the brain unchanged. In the fishes we find an essential and considerable modification of the five vesicles; it is clearly the brain of the Selachii in the first place, and subsequently the brain of the Ganoids, from which the brain of the rest of the fishes on the one hand and of the Dipneusts and Amphibia, and through these of the higher Vertebrates, on the other hand, must be derived. In the fishes and Amphibia (Fig. 300) there is a preponderant development of the middle brain, and also the after brain, the first, second, and fourth sections remaining very primitive. It is just the reverse in the higher Vertebrates, in which the first and third sections, the cerebrum and cerebellum, are exceptionally developed; while the middle brain and after brain remain small. The corpora quadrigemina are mostly covered by the cerebrum, and the oblongata by the cerebellum. But we find a number of stages of development within the higher Vertebrates themselves. From the Amphibia upwards the brain (and with it the psychic life) develops in two different directions; one of these is followed by the reptiles and birds, and the other by the mammals. The development of the first section, the fore brain, is particularly characteristic of the mammals. It is only in them that the cerebrum becomes so large as to cover all the other parts of the brain (Figs. 293, 301–304).
