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order. They resemble the round cell-communities of the Magospherae and Volvocina, equivalent to the ontogenetic blastula: hollow globules, the wall of which consists of a single layer of ciliated cells (blastoderm).

THIRD STAGE: THE GASTRAEADS.

Man's ancestors are Gastraeads, like the simplest of the actual Metazoa (Prophysema, Olynthus, Hydra, Pemmatodiscus). Their body consists merely of a primitive gut, the wall of which is made up of the two primary germinal layers.

FOURTH STAGE: THE PLATODES.

Man's ancestors have substantially the organisation of simple Platodes (at first like the cryptocoelic Platodaria, later like the rhabdocoelic Turbellaria). The leaf-shaped bilateral-symmetrical body has only one gut-opening, and develops the first trace of a nervous centre from the ectoderm in the middle line of the back (Figures 2.239 and 2.240).

FIFTH STAGE: THE VERMALIA.

Man's ancestors have substantially the organisation of unarticulated Vermalia, at first Gastrotricha (Ichthydina), afterwards Frontonia (Nemertina, Enteropneusta). Four secondary germinal layers develop, two middle layers arising between the limiting layers (coeloma). The dorsal ectoderm forms the vertical plate, acroganglion (Figure 2.243).

SIXTH STAGE: THE PROCHORDONIA.

Man's ancestors have substantially the organisation of a simple unarticulated Chordonium (Copelata and Ascidia-larvae). The unsegmented chorda develops between the dorsal medullary tube and the ventral gut-tube. The simple coelom-pouches divide by a frontal septum into two on each side; the dorsal pouch (episomite) forms a muscle-plate; the ventral pouch (hyposomite) forms a gonad. Head-gut with gill-clefts.

SEVENTH STAGE: THE ACRANIA.

Man's ancestors are skull-less Vertebrates, like the Amphioxus. The body is a series of metamera, as several of the primitive segments are developed. The head contains in the ventral half the branchial gut, the trunk the hepatic gut. The medullary tube is still simple. No skull, jaws, or limbs.

EIGHTH STAGE: THE CYCLOSTOMA.

Man's ancestors are jaw-less Craniotes (like the Myxinoida and Petromyzonta). The number of metamera increases. The fore-end of the medullary tube expands into a vesicle and forms the brain, which soon divides into five cerebral vesicles. In the sides of it appear the three higher sense-organs: nose, eyes, and auditory vesicles. No jaws, limbs, or floating bladder.

NINTH STAGE: THE ICHTHYODA.

Man's ancestors are fish-like Craniotes: (1) Primitive fishes (Selachii); (2) plated fishes (Ganoida); (3) amphibian fishes (Dipneusta); (4) mailed amphibia (Stegocephala). The ancestors of this series develop two pairs of limbs: a pair of fore (breast-fins) and of hind (belly-fins) legs. The gill-arches are formed between the gill-clefts: the first pair form the maxillary arches (the upper and lower jaws). The floating bladder (lung) and pancreas grow out of the gut.

TENTH STAGE: THE AMNIOTES.

Man's ancestors are Amniotes or gill-less Vertebrates: (1) Primitive Amniotes (Proreptilia); (2) Sauromammals; (3) Primitive Mammals (Monotremes); (4) Marsupials; (5) Lemurs (Prosimiae); (6) Western apes (Platyrrhinae); (7) Eastern apes (Catarrhinae): at first tailed Cynopitheca; then tail-less anthropoids; later speechless ape-men (Alali); finally speaking man. The ancestors of these Amniotes develop an amnion and allantois, and gradually assume the mammal, and finally the specifically human, form.

CHAPTER IX(24. EVOLUTION OF THE NERVOUS SYSTEM.)

 

The previous chapters have taught us how the human body as a whole develops from the first simple rudiment, a single layer of cells. The whole human race owes its origin, like the individual man, to a simple cell. The unicellular stem-form of the race is reproduced daily in the unicellular embryonic stage of the individual. We have now to consider in detail the evolution of the various parts that make up the human frame. I must, naturally, confine myself to the most general and principal outlines; to make a special study of the evolution of each organ and tissue is both beyond the scope of this work, and probably beyond the anatomic capacity of most of my readers to appreciate. In tracing the evolution of the various organs we shall follow the method that has hitherto guided us, except that we shall now have to consider the ontogeny and phylogeny of the organs together. We have seen, in studying the evolution of the body as a whole, that phylogeny casts a light over the darker paths of ontogeny, and that we should be almost unable to find our way in it without the aid of the former. We shall have the same experience in the study of the organs in detail, and I shall be compelled to give simultaneously their ontogenetic and phylogenetic origin. The more we go into the details of organic development, and the more closely we follow the rise of the various parts, the more we see the inseparable connection of embryology and stem-history. The ontogeny of the organs can only be understood in the light of their phylogeny, just as we found of the embryology of the whole body. Each embryonic form is determined by a corresponding stem-form. This is true of details as well as of the whole.

We will consider first the animal and then the vegetal systems of organs of the body. The first group consists of the psychic and the motor apparatus. To the former belong the skin, the nervous system, and the sense-organs. The motor apparatus is composed of the passive and the active organs of movement (the skeleton and the muscles). The second or vegetal group consists of the nutritive and the reproductive apparatus. To the nutritive apparatus belong the alimentary canal with all its appendages, the vascular system, and the renal (kidney) system. The reproductive apparatus comprises the different organs of sex (embryonic glands, sexual ducts, and copulative organs).

As we know from previous chapters (1.11 to 1.13), the animal systems of organs (the organs of sensation and presentation) develop for the most part out of the OUTER primary germ-layer, or the cutaneous (skin) layer. On the other hand, the vegetal systems of organs arise for the most part from the INNER primary germ-layer, the visceral layer. It is true that this antithesis of the animal and vegetal spheres of the body in man and all the higher animals is by no means rigid; several parts of the animal apparatus (for instance, the greater part of the muscles) are formed from cells that come originally from the entoderm; and a great part of the vegetative apparatus (for instance, the mouth-cavity and the gonoducts) are composed of cells that come from the ectoderm.

In the more advanced animal body there is so much interlacing and displacement of the various parts that it is often very difficult to indicate the sources of them. But, broadly speaking, we may take it as a positive and important fact that in man and the higher animals the chief part of the animal organs comes from the ectoderm, and the greater part of the vegetative organs from the entoderm. It was for this reason that Carl Ernst von Baer called the one the animal and the other the vegetative layer (see

Chapter 1.

3).

The solid foundation of this important thesis is the gastrula, the most instructive embryonic form in the animal world, which we still find in the same shape in the most diverse classes of animals. This form points demonstrably to a common stem-form of all the Metazoa, the Gastraea; in this long-extinct stem-form the whole body consisted throughout life of the two primary germinal layers, as is now the case temporarily in the gastrula; in the Gastraea the simple cutaneous (skin) layer ACTUALLY represented all the animal organs and functions, and the simple visceral (gut) layer all the vegetal organs and functions. This is the case with the modern Gastraeads (Figure 2.233); and it is also the case potentially with the gastrula.

We shall easily see that the gastraea theory is thus able to throw a good deal of light, both morphologically and physiologically, on some of the chief features of embryonic development, if we take up first the consideration of the chief element in the animal sphere, the psychic apparatus or sensorium and its evolution. This apparatus consists of two very different parts, which seem at first to have very little connection with each other--the outer skin, with all its hairs, nails, sweat-glands, etc., and the nervous system. The latter comprises the central nervous system (brain and spinal cord), the peripheral, cerebral, and spinal nerves, and the sense-organs. In the fully-formed vertebrate body these two chief elements of the sensorium lie far apart, the skin being external to, and the central nervous system in the very centre of, the body. The one is only connected with the other by a section of the peripheral nervous system and the sense-organs. Nevertheless, as we know from human embryology, the medullary tube is formed from the cutaneous layer. The organs that discharge the most advanced functions of the animal body--the organs of the soul, or of psychic life--develop from the external skin. This is a perfectly natural and necessary process. If we reflect on the historical evolution of the psychic and sensory functions, we are forced to conclude that the cells which accomplish them must originally have been located on the outer surface of the body. Only elementary organs in this superficial position could directly receive the influences of the environment. Afterwards, under the influence of natural selection, the cellular group in the skin which was specifically "sensitive" withdrew into the inner and more protected part of the body, and formed there the foundation of a central nervous organ. As a result of increased differentiation, the skin and the central nervous system became further and further separated, and in the end the two were only permanently connected by the afferent peripheral sensory nerves.

(FIGURE 2.284. The human skin in vertical section (from Ecker), highly magnified, a horny layer of the epidermis, b mucous layer of the epidermis, c papillae of the corium, d blood-vessels of same, ef ducts of the sweat-glands (g), h fat-glands in the corium, i nerve, passing into a tactile corpuscle above.)

The observations of the comparative anatomist are in complete accord with this view. He tells us that large numbers of the lower animals have no nervous system, though they exercise the functions of sensation and will like the higher animals. In the unicellular Protozoa, which do not form germinal layers, there is, of course, neither nervous system nor skin. But in the second division of the animal kingdom also, the Metazoa, there is at first no nervous system. Its functions are represented by the simple cell-layer of the ectoderm, which the lower Metazoa have inherited from the Gastraea (Figure 1.30 e). We find this in the lowest Zoophytes--the Gastraeads, Physemaria, and Sponges (Figures 2.233 to 2.238). The lowest Cnidaria (the hydroid polyps) also are little superior to the Gastraeads in structure. Their vegetative functions are accomplished by the simple visceral layer, and their animal functions by the simple cutaneous layer. In these cases the simple cell-layer of the ectoderm is at once skin, locomotive apparatus, and nervous system.

(FIGURE 2.285. Epidermic cells of a human embryo of two months. (From Kolliker.))

When we come to the higher Metazoa, in which the sensory functions and their organs are more advanced, we find a division of labour among the ectodermic cells. Groups of sensitive nerve cells separate from the ordinary epidermic cells; they retire into the more protected tissue of the mesodermic under-skin, and form special neural ganglia there. Even in the Platodes, especially the Turbellaria, we find an independent nervous system, which has separated from the outer skin. This is the "upper pharyngeal ganglion," or acroganglion,

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