The Origin of Vertebrates

by Walter Holbrook Gaskell

This is what Kowalewsky has done. Thus he states that Metschnikoff had fed Mysis with carmine-grains, and found tubules at the base of the thoracic feet coloured red with carmine. He himself used an allied species, Parapodopsis cornutum, and found here also that the carmine was taken up by tubules situated in the basal segments of the feet. In Nebalia, feeding experiments with alizarin blue and carmine stained the antennal glands, and showed the existence of glands at the base of the eight thoracic feet. These glands resemble the foot-glands of Mysis, Parapodopsis, and Palæmon, and lie in the space through which the blood passes from the thoracic feet, i.e. from the gills, to the heart. In Squilla also, in addition to the shell-glands, special glands were discovered on the branchial feet on the path of the blood to the heart. These glands form continuous masses of cells which constitute large compact glands at the base of the branchial feet. Single cells of the same sort are found along the whole course of the branchial venous canal, right up to the pericardium.

These observations show that the Crustacea possess not only true excretory organs in the shape of coxal glands, i.e. antennary glands, shell-glands, etc., in the cephalic region, but also a series of segmental glands situated at the base of the appendages, especially of the respiratory appendages: a system, that is to say, of coxal glands which have lost their excretory function, through having lost their external opening, but have not in consequence disappeared, but still remain in situ, and still retain an important excretory function, having become lymphatic glands containing leucocytes. Such glands are especially found in the branchial appendages, and are called branchial glands by Cuénot, who describes them for all Decapoda.

Further, it is significant that the same method reveals the existence in Pantopoda of a double set of glands of similar character, one set in the basal segments of the appendage, and the other in the adjacent part of the body.

In scorpions also, Kowalewsky has shown that the remarkable lymphatic organ situated along the whole length of the nerve-cord in the abdominal region takes up carmine grains and bacilli; an organ which in Androctonus does not form one continuous gland, but a number of separate, apparently irregularly grouped, glandular bodies.

In addition to this median lymphatic gland, Kowalewsky has discovered in the scorpion a pair of lateral glands, to which he gives the name of lymphoid glands, which communicate with the thoracic body-cavity (i.e. the pseudocœle), are phagocytic, and, according to him, give origin to leucocytes by the proliferation of their lining cells, thus, as he remarks, reminding us of the nephridial capsules of Clepsine. These glands are so closely related in position to the coxal glands on each side that he has often thought that the lumen of the gland communicated with that of the coxal gland; he, however, has persuaded himself that there is no true communication between the two glands. Neither of these organs appears to be segmental, and until we know how they are developed it is not possible to say whether they represent fused segmental organs or not.

The evidence, then, is very strong that in the Crustacea and Arachnida the original segmental excretory organs do not disappear, but remain as ductless glands, of the nature of lymphatic glands, which supply leucocytes to the system.

Further, the evidence shows that the nephric organs, or parts of the cœlom in close connection with these organs, may be transformed into ductless glands, which do not necessarily contain free leucocytes as do lymph-glands, but yet are of such great importance as excretory organs that their removal profoundly modifies the condition of the animal. Such a gland is the so-called adrenal or suprarenal body, disease of which is a feature of Addison's disease; a gland which forms and presumably passes into the blood a substance of remarkable power in causing contraction of blood-vessels, a substance which has lately been prepared in crystalline form by Jokichi Takamine, and called by him "adrenalin"; a gland, therefore, of very distinctly peculiar properties, which cannot be regarded as rudimentary, but is of vital importance for the due maintenance of the healthy state.

In the Elasmobranchs two separate glandular organs have been called suprarenal; a segmental series of paired organs, each of which possesses a branch from the aorta and a sympathetic ganglion, and an unpaired series in close connection with the kidneys, to which Balfour gave the name of interrenal glands. Of these two sets of glands, Swale Vincent has shown that the extract of the interrenals has no marked physiological effect, in this respect resembling the extract of the cortical part of the mammalian gland, while the extract of the paired segmental organs of the Elasmobranch produces the same remarkable rise of blood-pressure as the extract of the medullary portion of the mammalian gland.

The development also of these two sets of glands is asserted to be different. Balfour considered that the suprarenals were derived from sympathetic ganglion-cells, but left the origin of the interrenals doubtful. Weldon showed that the cortical part of the suprarenals in the lizard was derived from the wall of the glomerulus of a number of mesonephric tubules. In Pristiurus, he stated that the mesoblastic rudiment described by Balfour as giving origin to the interrenals is derived from a diverticulum of each segmental tubule, close to the narrowing of its funnel-shaped opening into the body-cavity. With respect to the paired suprarenals he was unable to speak positively, but doubted whether they were derived entirely from sympathetic ganglia.

Weldon sums up the results of his observations by saying: "That all vertebrates except Amphioxus have a portion of the kidney modified for some unknown purpose not connected with excretion; that in Cyclostomes the pronephros alone is so modified, in Teleostei the pro- and part of the meso-nephros; while in the Elasmobranchs and the higher vertebrates the mesonephros alone gives rise to this organ, which has also in these forms acquired a secondary connection with certain of the sympathetic ganglia."