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There are doubtless many other phenomena which, though commonly assigned to reflex stimulation, are really due to direct stimulation. Research might profitably be turned towards the elucidation of this point. Since there is demonstrable evidence that a nerve when no longer in connection with its centre, or with ganglionic cells, may be excited by electricity, pressure, thermal and chemical stimuli, we must conclude that even when it is in connection with its centre, any local irritation from pressure, changes in the circulation, etc., will also excite it. But as such local excitations will have only local and isolated effects, they will rarely be conspicuous.

CHAPTER VI.
WHAT IS TAUGHT BY EMBRYOLOGY?

97. Subject to the qualification expressed in the last chapter, stimulation of muscles and glands involves a neural process in ingoing nerve, centre, and outgoing nerve. These are the triple elements of the โ€œnervous arc.โ€ If muscles were directly exposed to external influences, they would be stimulated without the intervention of a centre; but as a matter of fact they never are thus exposed, being always protected by the skin. Did the skin-nerves pass directly to the muscles underneath, they would move those muscles, without the intervention of a centre; but as a matter of fact the skin-nerves pass directly to a centre, so that it is only through a centre that they can act upon the muscles. Were muscles and glands directly connected with sensitive surfaces, their activity would indeed be awakened by direct stimulation; but unless the muscles were so connected the one with the other, by anastomosis of fibres or continuity of tissue, that the movement of one was the movement of all, there would need to be some other channel by which their separate energies should be combined and co-ordinated. In the higher organisms anastomosis of muscles is rare, and the combination is effected by means of the nerves.

98. Although analysis distinguishes the two elements of the neuro-muscular system, assigning separate properties to the separate tissues, an interpretation of the phenomena demands a synthesis, so that a movement is to be conceived as always involving Sensibility, and a sensation as always involving Motility.126 In like manner, although analysis distinguishes the various organs of the body, assigning separate functions to each, our interpretation demands their synthesis into an organism; and we have thus to explain how the whole has different parts, and how these different parts are brought into unity. Embryology helps us to complete the fragmentary indications of Anatomy and Physiology.

99. Take a newly laid egg, weigh it carefully, then hatch it, and when the chick emerges, weigh both chick and shell: you will find that there has been no increase of weight. The semifluid contents have become transformed into bones, muscles, nerves, tendons, feathers, beak, and claws, all without increase of substance. There has been differentiation of structure, nothing else. Oxygen has passed into it from without; carbonic acid has passed out of it. The molecular agitation of heat has been required for the rearrangements of the substance. Without oxygen there would have been no development. Without heat there would have been none. Had the shell been varnished, so as to prevent the due exchange of oxygen and carbonic acid, no chick would have been evolved. Had only one part of the shell been varnished, the embryo would have been deformed.

99a. The patient labors of many observers (how patient only those can conceive who have made such observations!) have detected something of this wondrous history, and enabled the mind to picture some of the incessant separations and reunions, chemical and morphological. Each stage of evolution presents itself as the consequence of a preceding stage, at once an emergence and a continuance; so that no transposition of stages is possible; each has its appointed place in the series (Problem I. ยง 107). For in truth each stage is a processโ€”the sum of a variety of co-operant conditions. We, looking forward, can foresee in each what it will become, as we foresee the man in the lineaments of the infant; but in this prevision we always presuppose that the regular course of development will proceed unchecked through the regular succession of special conditions: the infant becomes a man only when this succession is uninterrupted. Obvious as this seems, it is often disregarded; and the old metaphysical conception of potential powers obscures the real significance of Epigenesis. The potentiality of the cells of the germinal membrane is simply their capability of reaching successive stages of development under a definite series of co-operant conditions. We foresee the result, and personify our prevision. But that result will not take place unless all the precise changes that are needful serially precede it. A slight pressure in one direction, insufficient to alter the chemical composition of the tissue, may so alter its structure as to disturb the regular succession of forms necessary to the perfect evolution.

100. The egg is at first a microscopic cell, the nucleus of which divides and subdivides as it grows. The egg becomes a hollow sphere, the boundary wall of which is a single layer of cells, all so similar that to any means of appreciation we now possess they are indistinguishable. They are all the progeny of the original nucleus and yolk, or cell contents. Very soon, however, they begin to show distinguishable differences, not perhaps in kind, but in degree. The wall of this hollow sphere is rapidly converted into the germinal membrane, out of which the embryo is formed. Kowalewsky (confirmed by Balfour) has pointed out how in the Amphioxus the hollow sphere first assumes an oval shape, and then, by an indentation of the under side, with corresponding curvature of the upper side, presents somewhat the shape of a bowl. The curvature increases, and the curved ends approaching each other, the original cavity is reduced to a thin line separating the upper from the under surface. The cavity of the body is formed by the curving downwards of this double layer of the germinal membrane.

101. This is not precisely the course observable in other vertebrates; but in all, the germinal membrane, which lies like a watch-glass on the surface of the yolk, is recognizable as two distinct layers of very similar cells. What do these represent? They are the starting-points of the two great systems: Instrumental and Alimental. The one yields the dermal surface; the other the mucous membrane. Each follows an independent though analogous career. The yolk furnishes nutrient material to the germinal membrane, and so passes more or less directly into the tissues; but unlike the germinal membrane, it is not itself to any great extent the seat of generation by segmentation. There are two yolks: the yellow and the white (which must not be confounded with what is called the white of egg); and their disposition may be seen in the diagram (Fig. 14) copied from Foster and Balfourโ€™s work. The importance of the white yolk is that it passes insensibly into a distinct layer of the germinal membrane, between the two primary layers.127 Each of the three layers of the germinal membrane has its specific character assigned to it by embryologists, who, however, are not all in agreement. Some authorities regard the topmost layer as the origin of the nervous system, the epidermis, with hair, feathers, nails, horns, the cornea and lens of the eye, etc. To the middle layer are assigned the muscular and osseous systems, the sexual organs, etc. To the innermost layer, the alimentary canal, with liver, pancreas, gastric and enteric glands. Other authorities are in favor of two primary layers: one for the nervous, muscular, osseous, and dermal systems; the other for the viscera and unstriped muscles. Between these two layers, a third gradually forms, which is specially characterized as the vascular.

Fig. 14.โ€”Diagrammatic section of an unincubated henโ€™s egg. bl, blastoderm; w y, white yolk; y y, yellow yolk; v t, vitelline membrane; x and w, layers of albumen; ch l, chalaza; a ch, air-chamber; i s m, internal layer of shell membrane; s m, external layer; s, shell.

102. Messrs. Foster and Balfour, avoiding the controverted designations of serous, vascular, and mucous layers, or of sensorial, motor germinative, and glandular layers, employ designations which are independent of theoretic interpretation, and simply describe the position of the layers, namely, epiblast for the upper, mesoblast for the middle, and hypoblast for the under layer. From the epiblast they derive the epidermis and central nervous system (or would even limit the latter to the central gray matter), together with some parts of the sense-organs. From the mesoblast, the muscles, nerves (and probably white matter of the centres), bones, connective tissue, and blood-vessels. From the hypoblast, the epithelial lining of the alimentary canal, trachea, bronchial tubes, as well as the liver, pancreas, etc.128 Kรถllikerโ€™s suggestion is much to the same effect, namely, that the three layers may be viewed as two epithelial layers, between which subsequently arises a third, the origin of nerves, muscles, bones, connective tissue, and vessels.129

103. The way in which the history may be epitomized is briefly this: There are two germinal membranes, respectively representing the Instrumental and Alimental Systems. Each membrane differentiates, by different appropriations of the yolk substance, into three primary layers, epithelial, neural, and muscular. In the epiblast, or upper membrane, these layers represent: 1ยฐ, the future epidermis with its derivativesโ€”hair, feathers, nails, skin glands, and chromatophores; 2ยฐ, the future nervous tissue; 3ยฐ, the future muscular tissue.130 (Bone, dermis, connective tissue, and blood-corpuscles are subsequent formations.)

The hypoblast, or under membrane, in an inverted order presents a similar arrangement: 1ยฐ, the unstriped muscular tissue of viscera and vessels; 2ยฐ, the nervous tissue of the sympathetic system; 3ยฐ, the epithelial lining of the alimentary canal with its glands.

Fundamentally alike as these two membranes are, they have specific differences; but in both we may represent to ourselves the embryological unit constituted by an epithelial cell, a nerve-cell, and a muscle-cell. All the other cells and tissues are adjuncts, necessary, indeed, to the working of the vital mechanism, but subordinated to the higher organites.

104. This conception may be compared with that of His in the division of Archiblast and Parablast assigned by him to the germ and accessory germ.131 We can imagine, he says, the whole of the connective substances removed from the organism, and thus leave behind a scaffolding in which brain and spinal cord would be the axis, surrounded by muscles, glands, and epithelium, and nerves as connecting threads. All these parts stand in more or less direct relation to the nervous system. All are continuous. By a similar abstraction we can imagine this organic system removed, and leave behind the connected scaffolding which is formed from the accessory germ; but this latter has only mechanical significance; the truly vital functions belong to the other system.

105. The researches of modern histologists have all converged towards the conclusion that the organs of Sense are modifications of the surface, with epithelial cells which on the one side are connected with terminal hairs, or other elements adapted to the reception of stimuli, and are connected on the other side through nerve-fibres with the perceptive centres. It has been shown that nerve-fibres often terminate in (or among) epithelial cellsโ€”sensory fibres at the surface, and motor-fibres in the glands.132 Whether the fibres actually penetrate the substance of the cell, or not, is still disputed. Enough for our present purpose to understand that there is a physiological connection between the two, and above all that sensory nerves are normally stimulated through some epithelial structure or other.

Fig. 15.โ€”Transverse

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