A, supra-œsophageal ganglia; B, infra-œsophageal ganglia; Al, œsophagus.

Truly, at the time when vertebrates first appeared, the direction and progress of variation in the Arthropoda was leading, owing to the manner in which the brain was pierced by the œsophagus, to a terrible dilemma—either the capacity for taking in food without sufficient intelligence to capture it, or intelligence sufficient to capture food and no power to consume it.

Something had to be done—some way had to be found out of this difficulty. The atrophy of the brain meant degeneration and the reduction to a lower stage of organization, as is seen in the Tunicata. The further development of the brain necessitated the establishment of a new method of alimentation and the closure of the old œsophagus, its vestiges still remaining as the infundibular canal of the vertebrate, meant the enormous upward stride of the formation of the vertebrate.

At first sight it might appear too great an assumption even to imagine the possibility of the formation of a new gut in an animal so highly organized as an arthropod, but a little consideration will, I think, show that such is not the case.

In the higher animals we are accustomed to speak of certain organs as vital and necessary for the further existence of the animal; these are essentially the central nervous system, the respiratory system, the circulatory system, and the digestive system. Of these four vital systems the first cannot be touched without the chance of degeneration; but that is not the case with the second. The passage from the fish to the amphibian, from the water-breathing to the air-breathing animal, has actually taken place, and was effected by the modification of the swim-bladder to form new respiratory organs—the lung; the old respiratory organs—the gills—becoming functionless, but still persisting in the embryo as vestiges. The necessity arose in consequence of the passage of the animal from water to land, and with this necessity nature found a means of overcoming the difficulty; air-breathing vertebrates arose, and from the very fact of their being able to extend over the land-surfaces, increased in numbers and developed in complexity in the manner already sketched out.

For a respiratory system all that is required is an arrangement by means of which blood should be brought to the surface, so as to interchange its gases with those of the external medium; and it is significant to find that of all vertebrates the Amphibia alone are capable of an effective respiration by means of the skin.

As to the circulatory system, it is exceedingly easily modified. An animal such as Amphioxus has no heart; in some the heart is systemic, in others branchial; in some there are more than one heart; in others there are contractile veins in addition to a heart. There is no difficulty here in altering and modifying the system according to the needs of the individual.

For a digestive system all that is required is an arrangement for the digestion and absorption of food, a mechanism which can arise easily if some of the cells of the skin possess digestive power. Now Miss Alcock has shown that some of the surface-cells of crustaceans secrete a fluid which possesses digestive powers, and she has also shown that certain of the cells in the skin of Ammocœtes possess digestive power.

The difficulty, then, of forming a new digestive system in the passage from the arthropod to the vertebrate is very much the same as the difficulty in forming a new respiratory system in the passage from the water-breathing fish to the air-breathing amphibian—a change which does not strike us as inconceivable, because we know it has taken place.

The whole argument so far leads to the conclusion that vertebrates arose from ancient forms of arthropods by the formation of a new alimentary canal, and the enclosure of the old canal by the growing central nervous system. If this conclusion is true, then it follows that we possess a well-defined starting-point from which to compare the separate organs of the arthropod with those of the vertebrate, and if, in consequence of such working hypothesis, each organ of the arthropod is found in the vertebrate in a corresponding position and of similar structure, then the truth of the starting-point is proved as fully as can possibly be expected by deductive methods. It is, in fact, this method of comparative anatomy which has proved the descent of man from the ape, the frog from the fish, etc.