In order to make clear the nature of the branchial segments in Ammocœtes, I have divided the head-part of the animal by means of a longitudinal horizontal section into halves—ventral and dorsal—as shown in Figs. 63 and 64. These figures are each a combination of a section and a solid drawing. The animal was slit open by a longitudinal section in the neighbourhood of the gill-slits, and each half was slightly flattened out, so as to expose the ventral and dorsal internal surfaces respectively. The structures in the cut surface were drawn from one of a series of horizontal longitudinal sections taken through the head of the animal. These figures show that the head-region of Ammocœtes consists of two chambers, the contents of which are different. In front, an oral or stomodæal chamber, which contains the velum and tentacles, is enclosed by the upper and lower lips, and was originally separated by a septum from the larger respiratory chamber, which contains the separate pairs of branchiæ. A glance at the two drawings shows clearly that Rathke's original description of this chamber is the natural one, for he at that time, looking upon Ammocœtes branchialis as a separate species, described the branchial chamber as containing a series of paired gills, with the gill-openings between consecutive gills. His branchial unit or gill, therefore, was represented by each of the so-called diaphragms, which, as seen in Figs. 63, 64, are all exactly alike, except the first and the last. Any one of these is represented in section in Fig. [65], and represents a branchial unit in Rathke's view and in mine. Clearly, it may be described as a branchial appendage which projects into an open pharyngeal chamber, so that the series of such appendages divides the chamber into a series of compartments, each of which communicates with the exterior by means of a gill-slit, and with each other by means of the open space between opposing appendages.
Each of these appendages possesses its own cartilaginous bar (Br. cart.), as explained in Chapter III.; each possesses its own branchial or visceral muscles (coloured blue in Figs. 63 and 64), separated absolutely from the longitudinal somatic muscles (coloured dark red in Figs. 63 and 64) by a space (Sp.) containing blood and peculiar fat-cells, etc. Each possesses its own afferent branchial blood-vessel from the ventral aorta, and its own efferent vessel to the dorsal aorta (Fig. [65], a. br. and v. br.). Each possesses its own segmental nerve, which supplies its own branchial muscles and no others with motor fibres, and sends sensory fibres to the general surface of each appendage, as also to the special sense-organs in the shape of the epithelial pits (S., Fig. [65]) arranged along the free edges of the diaphragms; each of these nerves possesses its own ganglion—the epibranchial ganglion.
Fig. 63.—Ventral half of Head-region of Ammocœtes.
Somatic muscles coloured red. Branchial and visceral muscles coloured blue. Tubular constrictor muscles distinguished from striated constrictor muscles by simple hatching. Tent., tentacles; Tent. m.c., muco-cartilage of tentacles; Vel. m.c., muco-cartilage of the velum; Hy. m.c., muco-cartilage of the hyoid segment; Ps. br., pseudo-branchial groove; Br. cart., branchial cartilages; Sp., space between somatic and splanchnic muscles; Th. op., orifice of thyroid; H., heart.
Fig. 64.—Dorsal half of Head-region of Ammocœtes.
Tr., trabeculæ; Pit., pituitary space; Inf., infundibulum; Ser., median serrated flange of velar folds.
| Fig. 65.—Section through Branchial Appendage of Ammocœtes. br. cart., branchial cartilage; v. br., branchial vein; a. br., branchial artery; b.s., blood-spaces; p., pigment; S., sense-organ; c., ciliated band; E., I., external and internal borders; m. add., adductor muscle; m.c.s., striated constrictor muscle; m.c.t., tubular constrictor muscle; m. and m.v., muscles of valve. | Fig. 66.—Section through Branchial Appendage of Limulus. br. cart., branchial cartilage; v.br., branchial vein; b.s., blood-spaces formed by branchial artery; P., pigment; m1, posterior entapophysio-branchial muscle; m2, anterior entapophysio-branchial muscle; m3, external branchial muscle. |
The work of Miss Alcock has shown that the segmental branchial nerve supplies solely and absolutely such an appendage or branchial segment, and does not supply any portion of the neighbouring branchial segments. The nerve-supply in Ammocœtes gives no countenance to the view that the original unit was a branchial pouch, the two sides of which each nerve supplied, but is strong evidence that the original unit was a branchial appendage, which was supplied by a single nerve with both motor and sensory fibres.