The Origin of Vertebrates

by Walter Holbrook Gaskell

2. A large capillary network (the glomus) at the distal end.

3. A peculiar tissue (the 'strittige Gewebe' of the Semon-Spengel controversy), which Spengel considers to be composed of the altered epithelium of pronephric tubules, while Semon looks on it as an amalgamation of glomeruli.

Maas is entirely on the side of Spengel, and shows that this peculiar tissue is actually formed by modified pronephric tubules, which become more and more lymphatic in character.

He says: "The pronephros consists of a number of nephric tubules, placed separately one behind the other, which were originally segmental in character, each one of which is supplied by a capillary network from a segmental branch of the aorta. The tubules begin with many mouths (dorso-lateral and medial-ventral) in the pericardial cavity; on their other blind end they have lost their original external opening, and there, in the cranial portion of the head-kidney, before they have joined together to form a collecting duct, they, together with the vascular network, are transformed into a peculiar adrenal-like tissue. The most posterior of the segmental capillary nets retain their original character, and are concentrated into the separate capillary mass known as the glomus."

Later on he says: "Further, the separate head-kidney is more and more removed in structure from an excretory organ in the ordinary sense. One cannot, however, speak of it as an organ becoming rudimentary; this is proved not only by the progressive transformation of its internal tissue into a tissue of a very definite character, but also by the cilia in its canals, and the steady increase in the number of its funnels. It appears, therefore, to be the conversion of an excretory organ into an organ for the transference of fluid out of the cœlom into a special tissue, i.e. into its blood-sinus; in other words, into an organ which must be classed as belonging to the lymph-system."

In exact correspondence with this transformation of a nephric tubule into a ductless gland of the nature of a lymphatic gland, is the formation of the head-kidney in the Teleostea. Thus, Weldon points out that, though the observations of Balfour left it highly probable that the "lymphatic" tissue described by him was really a result of the transformation of part of the embryonic kidney, he did not investigate the details of its development. This was afterwards done by Emery, with the following results: "In those Teleostea which he has studied, Professor Emery finds that at an early stage the kidney consists entirely of a single pronephric funnel, opening into the pericardium, and connected with the segmental duct, which already opens to the exterior. Behind this funnel, the segmental duct is surrounded by a blastema, derived from the intermediate cell-mass, which afterwards arranges itself more or less completely into a series of solid cords, attaching themselves to the duct. These develop a lumen, and become normal segmental tubules, but it is, if I may be allowed the expression, a matter of chance how much of the blastema becomes so transformed into kidney tubules, and how much is left as the 'lymphatic' tissue of Balfour, this 'lymphatic' tissue remaining either in the pronephros only, or in both pro- and meso-nephros."

If we turn now to the invertebrates, we see also how close a connection exists between lymphatic and phagocytic organs and excretory organs. The chief merit for this discovery is due to Kowalewsky, who, taking a hint from Heidenhain's work on the kidney, in which he showed how easy it was to find out the nature of different parts of the mammalian excretory organ by the injection of different substances, such as a solution of ammoniated carmine, or of indigo-carmine, has injected into a large number of different invertebrates various colouring matters, or litmus, or bacilli, and thus shown the existence, not only of known excretory organs, but also of others, lymphatic or lymphoid in nature, not hitherto suspected.

In all cases he finds that a phagocytic action with respect to solid bodies is a property of the leucocytes, and that these leucocytes which are found in the cœlomic spaces of the Annelida, etc., are apparently derived from the epithelium of such spaces. Also by the proliferation of such epithelium in places, e.g. the septal glands of the terrestrial Oligochæta, segmental glandular masses of such tissue are formed which take up the colouring matter, or the bacilli. In the limicolous Oligochæta such septal glands are not found, but at the commencement of the nephridial organ, immediately following upon the funnel, a remarkable modification of the nephridial wall takes place to form a large cellular cavernous mass, the so-called filter, which in Euaxes is full of leucocytes; the cells are only definable by their nuclei, and look like and act in the same way as the free leucocytes outside this nephridial appendage. As G. Schneider points out, the whole arrangement is very like that described by Kowalewsky in the leeches Clepsine and Nephelis, where, also immediately succeeding the funnel of the nephridial organ, a large accessory organ is found, which is part of the nephridium, and is called the nephridial capsule. This is the organ par excellence which takes up the solid carmine-grains and bacilli, and apparently, from Kowalewsky's description, contains leucocytes in large numbers. We see, then, that in such invertebrates, just as in the vertebrate, modifications of the true excretory organ may give rise to phagocytic glands of the nature of lymphatic glands. Further, these researches of Kowalewsky suggest in the very strongest manner that whenever by such means new, hitherto unsuspected glands are discovered, such glands must belong to the excretory system, i.e. must be derived from cœlomic epithelium, even when all evidence of any cœlom has disappeared. Kowalewsky himself was evidently so impressed with the same feeling that he heads one of his papers "The Excretory Organs of the Pantopoda," although the organs in question had been discovered by him by this method, and appeared as ductless glands with no external opening.

To my mind these observations of Kowalewsky are of exceeding interest, for it is immediately clear that if the segmental organs of the annelids, which must have existed on all the segments of the forefathers of the Crustacea and Arachnida (the Protostraca), have left any sign of their existence in living crustaceans and arachnids, then such indication would most likely take the form of lymphatic glands in the places where the excretory organs ought to have been.

Now, as already pointed out in Peripatus, such segmental organs were formed by the ventral part of the cœlom, and dipped originally into each appendage. We know also that each segment of an arachnid embryo possesses a cœlomic cavity in its ventral part which extends into the appendage on each side; this cavity afterwards disappears, and is said to leave no trace in the adult of any excretory coxal gland derived from its walls. If, however, it is found that in the very position where such organ ought to have been formed a segmentally arranged ductless gland is situated, the existence of which is shown by its taking up carmine, etc., then it seems to me that in all probability such gland is the modification of the original coxal gland.