The whole structure is clearly remarkably like Rohon's picture of a section of the head-plate of a Cephalaspid (Fig. [130]). In the latter case the matrix contains calcium salts, in the former it is composed of the peculiar homogeneous mucoid tissue which stains so characteristically with thionin. With respect to this calcification, it is instructive to recall the calcification in the interior of the branchial cartilages of Limulus, as described in Chapter III., for this example shows how easy it is to obtain a calcification in this chondro-mucoid material. With respect to the medullary spaces and smaller spaces in this tissue, as described by Rohon, I would venture to suggest that they need not all necessarily indicate blood-vessels, for similar spaces would appear in the head-shield of Ammocœtes if its muco-cartilage alone were preserved. Of these, some would indicate the position of blood-vessels, such, for instance, as of the external carotid which traverses this structure; but the largest and most internal spaces, resembling Rohon's medullary spaces, would represent muscles, being filled up with bundles of the upper lip-muscles.
The Muco-Cartilaginous Head-Shield of Ammocœtes.
The resemblance between the structure of the head-shield of Thyestes and the muco-cartilage of Ammocœtes, is most valuable, for muco-cartilage is unique, occurs in no other vertebrate, and every trace of it vanishes at transformation; it is essentially a characteristic of the larval form, and must, therefore, in accordance with all that has gone before, be the remnant of an ancestral skeletal tissue. The whole story deduced from the study of Ammocœtes would be incomplete without some idea of the meaning of this tissue. So also, as already mentioned, the skeleton of Ammocœtes is incomplete without taking this tissue into account. It is confined entirely to the head-region; no trace of it exists posteriorly to the branchial basket-work. It consists essentially of dorsal and ventral head-shields, connected together by the tentacular, metastomal, and thyroid bars, as already described. The ventral shield forms the muco-cartilaginous plate of the lower lip and the plate over the thyroid gland, so that the skeleton ventrally is represented by Fig. [118], B, which shows how the cartilaginous bars of the branchial basket-work are separated from each other by this thyroid plate. At transformation, with the disappearance of this muco-cartilaginous plate, the bars come together in the middle line, as in the more posterior portion of the branchial basket-work.
The dorsal head-shield of muco-cartilage covers over the upper lip, sends a median prolongation over the median pineal eyes and a lateral prolongation on each side as far as the auditory capsules, giving the shape of the head-shield of muco-cartilage, as in Fig. 118, C.
Not only then is the structure of the head-shield of a Cephalaspid remarkably like the muco-cartilage of Ammocœtes, but also its general distribution strangely resembles that of the Ammocœtes muco-cartilage.
Now, these head-shields in the Cephalaspidæ and Tremataspidæ vary very much in shape, as is seen by the comparison of Tremataspis and Auchenaspis with Cephalaspis and Eukeraspis, and yet, undoubtedly, all these forms belong to a single group, the Osteostraci.
The conception that Ammocœtes is the solitary living form allied to this group affords a clue to the meaning of this variation of shape, which appears to me to be possible, if not indeed probable. There is a certain amount of evidence given in the development of Ammocœtes which indicates that the branchial region of its ancestors was covered with plates of muco-cartilage as well as the prosomatic region.
The evidence is as follows:—
The somatic muscles of Ammocœtes form a continuous longitudinal sheet of muscles along the length of the body, which are divided up by connective tissue bands into a series of imperfect segments or myotomes. This simple muscular sheet can be dissected off along the whole of the head-region of the animal, with the exception of the most anterior part, without interfering with the attachments or arrangements of the splanchnic muscular system in the least. The reason why this separation can be so easily effected is to be found in the fact that the two sets of muscles are not attached to the same fascia. The sheet of fascia to which the somatic muscles are attached is separated from the fascia which encloses the branchial cavity by a space (cf. Figs. 63 and 64) filled with blood-spaces and cells containing fat, in which space is also situated the cartilaginous branchial basket-work. These branchial bars are closely connected with the branchial sheet of fascia, and have no connection with the somatic fascia, their perichondrium forming part of the former sheet. Upon examination, this space is seen to be mainly vascular, the blood-spaces being large and frequently marked with pigment; but it also possesses a tissue of its own, recognized as fat-tissue by all observers. The peculiarity of the cells of this tissue is their arrangement; they are elongated cells arranged at right angles to the plates of fascia, just as the fibres of the muco-cartilage are largely arranged at right angles to their limiting plates of perichondrium. These cells do not necessarily contain fat; and when they do, the fat is found in the centre of each cell, and does not push the protoplasm of the cell to the periphery, as in ordinary fat cells.
In Fig. [132], B, I give a specimen of this tissue stained by osmic acid; in Fig. [132], A, I give a drawing of ordinary muco-cartilage taken from the plate of the lower lip; and in Fig. [133], A, a modification of the muco-cartilage taken from the velum, which shows the formation of a tissue intermediate between ordinary muco-cartilage and this branchial fat-tissue.