The Origin of Vertebrates

by Walter Holbrook Gaskell

The brain in these animals may be spoken of as composed of three parts—(1) the fused supra-œsophageal ganglia, (2) the fused prosomatic ganglia, and (3) the fused mesosomatic ganglia. Such a brain is strictly homologous with the vertebrate brain, which also is built up of three parts—(1) the part in front of the notochord, the prechordal or supra-infundibular brain, which consists of the cerebral hemispheres, together with the basal and optic ganglia and corresponds, therefore, to the supra-œsophageal mass, with its olfactory and optic divisions lying in front of the œsophagus; (2 and 3) the epichordal brain, composed of (2) a trigeminal and (3) a vagus division, of which the first corresponds strictly to the fused prosomatic ganglia, and the second to the fused mesosomatic ganglia. Further, just as in the embryo of an arthropod it is possible, with more or less accuracy, to see the number of neuromeres or original ganglia which have fused to form the supra- and infra-œsophageal portions of its brain, so also in the embryo of a vertebrate we are able at an early stage to gain an indication, more or less accurate, of the number of neuromeres which have built up the vertebrate brain. The further consideration of these neuromeres, and the evidence they afford as to the number of the prosomatic and mesosomatic ganglia which have formed the epichordal part of the vertebrate brain, must be left to the chapter on the segmentation of the cranial nerves.

The further continuation of this process of concentration of separate segments, together with the fusion of the nervous system with the tube of the alimentary canal, leads in the simplest manner to the formation of the spinal cord of the vertebrate from the metasomatic ganglia of the ventral chain of the arthropod.

The Antagonism between Cephalization and Alimentation.

This concentration of the nervous system in the head-region, together with an actual increase in the bulk of the cephalic nervous masses, constitutes the great principle upon which the law of upward progress or evolution in the animal kingdom is based, and it illustrates in a striking manner the blind way in which natural selection works; for, as already explained, the central nervous system arose as a ring round the mouth, in consequence of which, with the progressive evolution of the animal kingdom, the œsophagus necessarily pierced the central nervous system at the cephalic end. At the same time, the very fact that the evolution was progressive necessitated the concentration and increase of the nervous masses in this very same œsophageal region.

Progress on these lines must result in a crisis, owing to the inevitable squeezing out of the food-channel by the increasing nerve-mass; and, indeed, the fact that such a crisis had in all probability arisen at the time when vertebrates first appeared is apparent when we examine the conditions at the present time.

Those invertebrates whose central nervous system is most concentrated at the cephalic end belong to the arachnid group, among which are included the various living scorpion-like animals, such as Thelyphonus, Androctonus, etc.

As already mentioned, the giants of the Palæostracan age were Pterygotus, Slimonia, etc., all animals of the scorpion-type—in fact, sea-scorpions. Now, all these animals, spiders and scorpions, without exception, are blood-suckers, and in all of them the concentrated cephalic mass of nervous material surrounds an œsophagus the calibre of which is so small that nothing but a fluid pabulum can be taken into the alimentary canal; and even for that purpose a special suctorial apparatus has in some species been formed on the gastric side of the œsophagus for the purpose of drawing blood through this exceedingly narrow tube.

In Fig. [25] this increasing antagonism between brain-power and alimentation, as we pass from such a form as Branchipus to the scorpion, is illustrated, and in Fig. [26] the relative sizes of the œsophagus and the brain-mass surrounding it is shown. The section shows that the food channel is surrounded by the white and grey matter of the brain as completely as the central canal of the spinal cord of the vertebrate is surrounded by the white and grey nervous material.

Fig. 26.—Transverse Section through the Brain of a Young Thelyphonus.