The Origin of Vertebrates

by Walter Holbrook Gaskell

The origin of the retina from an optic epidermal plate in vertebrates, as in all other animals, brings the cephalic eyes of all animals into the same category, and leaves the vertebrate eye no longer in an isolated and unnatural position. In one point the retina of the vertebrate eye differs from that of a compound retina of an invertebrate; in the former, a striking supporting tissue exists, known as Müller's fibres, which is absent in the latter. This difference of structure is closely associated with another of the same character as in the central nervous system, viz. the apparent development of the nervous part from a tube. We see, in fact, that the retinal and nervous arrangements of the vertebrate eye are comparable with those of the arthropod eye, in precisely the same way and to the same extent as the nervous matter of the brain of the vertebrate is comparable with the brain of the arthropod. In both cases the nervous matter is, in structure, position, and function, absolutely homologous; in both cases there is found in the vertebrate something extra which is not found in the invertebrate—viz. a hollow tube, the walls of which, in the case of the brain, are utilized as supporting tissues for the nerve structures. The explanation of this difference in the case of the brain is the fundamental idea of my whole theory, namely, that the hollow tube is in reality the cephalic stomach of the invertebrate, around which the nervous brain-matter was originally grouped in precisely the same manner as in the invertebrate. What, then, are the optic diverticula?

"The formation of the eye," as taught by Balfour, "commences with the appearance of a pair of hollow outgrowths from the anterior cerebral vesicle. These outgrowths, known as the optic vesicles, at first open freely into the cavity of the anterior cerebral vesicle. From this they soon, however, become partially constricted, and form vesicles united to the base of the brain by comparatively narrow, hollow stalks, the rudiments of the optic nerves."

"After the establishment of the optic nerves, there takes place (1) the formation of the lens, and (2) the formation of the optic cup from the walls of the primary optic vesicle."

He then goes on to explain how the formation of the lens forms the optic cup with its double walls from the primary optic vesicle, and says—

"Of its double walls, the inner, or anterior, is formed from the front portion, the outer, or posterior, from the hind portion of the wall of the primary optic vesicle. The inner, or anterior, which very speedily becomes thicker than the other, is converted into the retina; in the outer, or posterior, which remains thin, pigment is eventually deposited, and it ultimately becomes the tesselated pigment-layer of the choroid."

The difficulties in connection with this view of the origin of the eye are exceedingly great, so great as to have caused Balfour to discuss seriously Lankester's suggestion that the eye must have been at one time within the brain, and that the ancestor of the vertebrate was therefore a transparent animal, so that light might get to the eye through the outer covering and the brain-mass; a suggestion, the unsatisfactory nature of which Balfour himself confessed. Is there really evidence of any part of either retina or optic nerve being formed from the epithelial lining of the tube?

This tube is formed as a direct continuation of the tube of the central nervous system, and we can therefore apply to it the same arguments as have been used in the discussion of the meaning of the latter tube. Now, the striking point in the latter case is the fact that the lining membrane of the central canal is in so many parts absolutely free from nervous matter, and so shows, as in the so-called choroid plexuses, its simple, non-nervous epithelial structure. This also we find in the optic diverticulum. Where there is no evidence of any invasion of the tube by nervous elements, there it retains its simple non-nervous character of a tube composed of a single layer of epithelial cells—viz. in that part of the tube which, as Balfour says, remains thin, in which pigment is eventually deposited, and which ultimately becomes the tesselated pigment-layer of the choroid. Nobody has ever suggested that this pigment-layer is nervous matter, or ever was, or ever will be, nervous matter; it is in precisely the same category as the membranous roof of the brain in Ammocœtes, which never was, and never will be, nervous matter. Yet, according to the old embryology both in the case of the eye and the brain, the pigment-layer and the so-called choroid plexuses are a part of the tubular nervous system.

Turning now to the optic nerve, Balfour describes it as derived from the hollow stalk of the optic vesicle. He says—

"At first the optic nerve is equally continuous with both walls of the optic cup, as must of necessity be the case, since the interval which primarily exists between the two walls is continuous with the cavity of the stalk. When the cavity within the optic nerve vanishes, and the fibres of the optic nerve appear, all connection is ruptured between the outer wall of the optic cup and the optic nerve, and the optic nerve simply perforates the outer wall, and becomes continuous with the inner one."

In this description Balfour, because he derived the optic nerve fibres from the epithelial wall of the optic stalk, of necessity supposed that such fibres originally supplied both the outer and inner walls of the optic cup and, therefore, seeing that when the fibres of the optic nerve appear they do not supply the outer wall, he supposes that their original connection with the outer wall is ruptured, because a discontinuity of the epithelial lining takes place coincidently with the appearance of the optic nerve-fibres, and, according to him, the optic nerve simply perforates the outer wall and becomes continuous with the inner one. This last statement is very difficult to understand. I presume he meant that some of the fibres of the optic nerve supplied from the beginning the inner wall of the optic cup, but that others which originally supplied the outer wall were first ruptured, then perforated the outer wall, and finally completed the supply to the inner wall or retina.