The Origin of Vertebrates

by Walter Holbrook Gaskell

Such an arrangement would lead directly to the condition found in Ammocœtes, if the generative material around the brain lost its function, owing to a new exit for generative products being formed in the posterior part of the body. The connection of the genital duct with this cephalic gland being then closed and cut off by the brain-case, the position of the oviducts would still be shown by the ciliated grooves opening into the folded-down thyroid tube, i.e. the folded-down horns of the uterus; the uterus itself would remain as the main body of the thyroid and still open by a conspicuous orifice into the common respiratory chamber. Next, in the degeneration process, we may suppose that not only the oviducts opened out to form the ciliated groove, but that the uterine chamber itself also opened out, and thus formed the endostyle of Amphioxus and of the Tunicata.

It might seem at first sight improbable that a closed tube should become an open groove, although the reverse phenomenon is common enough; the difficulty, however, is clearly not considered great, for it is precisely what Dohrn imagines to have taken place in the conversion of the thyroid of Ammocœtes into the endostyle of Amphioxus and the Tunicata; it is only carrying on the same idea a stage further to see in the open, ciliated groove of Ammocœtes the remains of the closed genital duct of Limulus and its allies.

Such is the conclusion to which the study of the thyroid gland in Ammocœtes seems to me to lead, and one cannot help wondering why such an unused and rudimentary organ should have remained after its original function had gone. Is it possible to find out its function in Ammocœtes?

The Function of the Thyroid Gland in Ammocœtes.

The thyroid gland has been supposed to secrete mucus into the respiratory chamber for the purpose of entangling the particles of food, and so aiding in digestion. I see no sign of any such function; neither by the thionin method, nor by any other test, have Miss Alcock and myself ever been able to see any trace of mucous secretion in the thyroid, and, indeed, the thyroid duct is always remarkably free from any sign of any secretion whatever. Not only is there no evidence of any mucous secretion in the thyroid of the fully developed Ammocœtes, but also no necessity for such secretion from Dohrn's point of view, for so copious a supply of mucus is poured out by the glands of the branchiæ, along the whole pharyngeal tract, especially from the cells of the foremost or hyoid gills, as to mix up with the food as thoroughly as can possibly be needed. Further, too, the ciliated pharyngeal bands described by Schneider are amply sufficient to move this mixed mass along in the way required by Dohrn. Finally, the evidence given by Miss Alcock is absolutely against the view that the thyroid takes any part in the process of digestion, while, on the other hand, her evidence directly favours the view that these glandular branchial mucus-secreting cells play a most important part in the digestive process.

In Fig. [93], A is a representation of the respiratory tissue of a normal gill; B is the corresponding portion of the first or hyoid gill, in which, as is seen, the whole of the respiratory epithelium is converted into gland-tissue of the nature of mucous cells.

To sum up, the evidence is clear and conclusive that the Ammocœtes possesses in its pharyngeal chamber mucus-secreting glands, which take an active part in the digestive process, which do not in the least resemble either in structure or arrangement the remarkable cells of the thyroid gland, and that the experimental evidence that the latter cells either secrete mucus or take any part in digestion is so far absolutely negative. It is, of course, possible, that they may contain mucin in the younger developmental stages, and therefore possible that they might at that stage secrete it; they certainly, however, show no sign of doing so in their more adult condition, and cannot be compared in the very faintest degree to the glandular cells of the pharyngeal region. It is also perfectly possible for gland-cells belonging to a retrograde organ to become mucus-secreting, and so to give rise to the cells of Amphioxus and the Tunicata.

Fig. 93.—A, Portion of a Gill of Ammocœtes with ordinary Respiratory Epithelium; B, Corresponding Portion of the First or Hyoid Gill.

If, then, these cells were not retained for digestive purposes, what was their function? To answer this question we must first know the function of the corresponding gland-cells in the uterus of the scorpion, which undoubtedly secreted into the cavity of the uterus and took some part in connection with the generative act, and certainly not with digestion. What the function of these cells is or in what way they act I am unable at present to say. I can only suppose that the reason why the thyroid gland has persisted throughout the vertebrate kingdom, after the generative tissues had found a new outlet for their products in the body-cavity of the posterior region, is because it possessed some important function in addition to that connected with the exit of the products of the generative organs; a function which was essential to the well-being, or even to the life of the animal. We do not know its function in the scorpion, or the nature of its secretion in that animal. We know only that physiology at the present day has demonstrated clearly that the actual external secretion of a gland may be by no means its most important function; in addition, glands possess what is called an internal secretion, viz. a secretion into the blood and lymph, and this latter secretion may be of the most vital importance. Now, the striking fact forces itself prominently forward, that the thyroid gland of the higher vertebrates is the most conspicuous example of the importance of such internal secretion. Here, although ductless, we have a gland which cannot be removed without fatal consequences. Here, in the importance of its internal secretion, we have a reason for the continued existence of this organ; an organ which remains much the same throughout the Vertebrata down to and including Petromyzon, but, as is seen at transformation, is all that remains of the more elaborate, more extensive organ of Ammocœtes. Surely we may argue that it is this second function which has led to the persistence of the thyroid, and that its original form, without its original function, is seen in Ammocœtes, because that is a larval form, and not a fully-developed animal. As soon as the generative organs of Petromyzon are developed at transformation, all trace of its connection with a genital duct vanishes, and presumably its internal secretory function alone remains.