Form and Function: A Contribution to the History of Animal Morphology

by E. S. Russell

Through morphological differentiation the various parts of the fundamental organs become specialised, through unequal growth, first into the primitive organs and then into the functional organs of the body. "Single sections of the tubes originally formed from the layers develop individual forms, which later acquire special functions: these functions are in the most general way subordinate elements of the function of the whole tube, but yet differ from the functions of other sections. Thus the nerve-tube differentiates into sense-organs, brain and spinal cord, the alimentary tube into mouth cavity, œsophagus, stomach, intestine, respiratory apparatus, liver, bladder, etc. This specialisation in development is bound up with increased or diminished growth" (p. 155). Rapid growth concentrated at one point brings about an evagination; in this manner are formed the sense-organs from the nerve-tube, the liver and lungs from the alimentary tube. Or increased growth over a section of a tube causes it to swell out; in this wise the brain develops from the nerve-tube, the stomach from the alimentary tube. The segmentation which soon becomes so marked, particularly in the muscle layer, is also due to a process of morphological differentiation.

At the same time that the organs of the body are being thus roughly blocked out and moulded from the germ-layers the third process of differentiation is actively going on. "In addition to the differentiation of the layers, there follows later another differentiation in the substance of the layers, whereby cartilage, muscle and nerve separate out, a part also of the mass becoming fluid and entering the bloodstream" (p. 154). Through histological differentiation the texture of the layers and incipient organs becomes individualised. In its earliest appearance the germ consists of an almost homogeneous mass, containing clear or dark globules suspended in its substance (ii., p. 92). This homogeneity gives place to heterogeneity; the structureless mass becomes fibrous to form muscles, hardens to form cartilage or bone, becomes liquid to form the blood, differentiates in a hundred other ways—into absorbing and secreting tissues, into nerves and ganglia, and so forth. It will be noticed that the concept of histological differentiation is independent of the cell-theory; it signifies that textural differentiation which leads to the formation of tissues in Bichat's sense. The tissues and the germ-layers stand in fairly close relation with one another, for while certain tissues are formed chiefly but not exclusively in one layer, others are formed only in one layer and never elsewhere. For example, peripheral nerves are for the most part formed in the muscle layer, though the bulk of the nervous tissue is formed in the walls of the nerve tube; similarly blood and blood-vessels may arise from almost any layer, though their chief seat of origin is the vessel-layer; on the other hand, bone is formed only in the muscle-layer (i., p. 155, ii., pp. 92-3).

This relation of tissue to germ-layer was more fully discussed and brought into greater prominence by Remak, from the standpoint of the cell-theory, and it will occupy us in a later chapter ([Chap. XII.]).

The fourth Scholion elaborates the analysis of developmental processes still further, and discusses in particular the scheme of development which is shown by the Vertebrata. The characteristic structure of the vertebrate body is brought about by a "double symmetrical" rolling together of the germ-layers, whereby two main tubes are formed, one above and one below the axis of the body, which is the chorda. The dorsal tube is formed by the two animal layers, the ventral tube by all the layers combined (see [Fig. 7]).

The process is indicated with sufficient clearness in the diagram. It will be seen that the real foundation and framework of the arrangement is the muscle-layer, with its two tubes, one surrounding the central nervous system and forming the "dorsal plates," the other surrounding the body cavity and forming the "ventral plates." In the dorsal plates, which early show metameric segmentation, the investing skeleton of the neural axis develops; in the ventral plates are formed the ribs, the ventral arches of the vertebræ, the hyoid, the lower jaw and other skeletal structures.

The alimentary or "mucous" tube and the part of the vessel layer which invests it become so closely bound up with one another as to form a single primitive organ—the alimentary canal. The muscles of the alimentary canal are accordingly in all probability developed in the investing part of the vessel layer. From the "Gekröse," or remaining part of the vessel layer develop the Wolffian bodies (Urnieren, Pronephros), the kidneys, the sex glands, and the series of "blood-glands"—suprarenals, thyroid, thymus and spleen. Baer did not attach any special morphological significance to the peritoneal lining of the body cavity, as is done in more modern forms of the germ-layer theory. The gill-slits were largely formed by outgrowths from the alimentary canal.

Fig. 7.—Ideal Transverse Section of a Vertebrate Embryo. (After von Baer.)

a. Chorda.	e. Vessel-layer.	i. Amnion.
b. Dorsal plates.	f. Alimentary tube.	k. Serous membrane.
c. Ventral plates.	g. Pronephros.	l. Tolk sac.
d. Spinal cord.	h. Skin.

In his germ-layer theory von Baer was influenced a good deal by Pander, to whom the actual discovery of the process of layer-formation is due. Pander, however, had distinguished only three germ-layers, an upper "serous" layer, a lower "mucous" layer and a middle "vessel-layer." He it was who introduced the terms "Keimhaut" (blastoderm) and "Keimblatt" (germ-layer).