Page:EB1911 - Volume 01.djvu/940

 lobe and the mouth. As already stated, the notochord extends beyond the mouth to the tip of the rostrum. The mouth consists of two portions, an outer vestibule and an inner apertura oris; the latter is surrounded by a sphincter muscle, which forms the so-called velum. The vestibule of the mouth is the space bounded by the oral hood; this arises by secondary downgrowth of lid-like folds over the true oral aperture, and is provided with a fringe of tentacular cirri, each of which is supported by a solid skeletal axis. The oral hood with its cirri has a special nerve supply and musculature by which the cirri can be either spread out, or bent inwards so that those of one side may interdigitate with those of the other, thus completely closing the entrance to the mouth. The velum is also provided with a circlet of twelve tentacles (in some species sixteen) which hang backwards into the pharynx; these are the velar tentacles. The atrial region extends from the mouth over about two-thirds of the length of the body, terminating at a large median ventral aperture, the atriopore; this is the excurrent orifice for the respiratory current of water and also serves for the evacuation of the generative products. This region is really the branchiogenital region, although the fact is not apparent in external view. The ventral side of the body in the atrial region is broad and convex, so that the body presents the appearance of a spherical triangle in transverse section, the apex being formed by the dorsal fin and the angles bordered by two hollow folds, the metapleural folds, each of which contains a continuous longitudinal lymph-space, the metapleural canal. In the genus Branchiostoma the metapleural folds terminate symmetrically shortly behind the atriopore, but in Heteropleuron the right metapleur passes uninterruptedly into the median crest of the ventral fin (fig. 1). In this connexion it may also be mentioned that in all cases the right half of the oral hood is directly continuous with the rostral fin (fig. 2). The abdominal region comprises a short stretch of body between atriopore and anus, the termination of the alimentary canal. It is characterized by the presence of a special development of the lophioderm or median fin-system, namely, the ventral fin, which is composed of two portions, a lower keel-like portion, which underlies an upper chambered portion, each chamber containing typically a pair of gelatinous fin rays. Finally, the caudal region comprises the post-anal division of the trunk. The keel of the ventral fin is continued past the anus into the expanded caudal fin, and so it happens that the anal opening is displaced from the middle line to the left side of the fin. In Asymmetron the caudal region is remarkable for the curious elongation of the notochord, which is produced far beyond the last of the myotomes.

Alimentary, Respiratory and Excretory Systems.—Although the function of the two latter systems of organs is the purification of the blood, they are not usually considered together, and it is therefore the more remarkable that their close association in Amphioxus renders it necessary to treat them in common. The alimentary canal is a perfectly straight tube lined throughout by ciliated epithelium. As food particles pass in through the mouth they become enveloped in a slimy substance (secreted by the endostyle) and conveyed down the gut by the action of the vibratile cilia as a continuous food-rope, the peristaltic movements of the gut-wall being very feeble. The first part of the alimentary canal consists of the pharynx or branchial sac, the side walls of which are perforated by upwards of sixty pairs of elongated slits, the gill-clefts. Each primary gill-cleft becomes divided into two by a tongue-bar which grows down secondarily from the upper wall of the cleft and fuses with the ventral wall. New clefts continue to form at the posterior end of the pharynx during the adult life of the animal. The gill-clefts open directly from the cavity of the pharynx into that of the atrium, and so give egress to the respiratory current which enters the mouth with the food (fig. 4). The atrium or atrial chamber is a peripharyngeal cavity of secondary origin effecting the enclosure of the gill-clefts, which in the larva opened directly to the exterior. The atrium is thus analogous to the opercular cavity of fishes and tadpoles, and, as stated above, remains in communication with the exterior by means of the atriopore. The primary and secondary bars which separate and divide the successive gill-clefts from one another are traversed by blood-vessels which run from a simple tubular contractile ventral branchial vessel along the bars into a dorsal aorta. The ventral branchial vessel lies below the hypobranchial groove or endostyle, and is the representative of a heart. As water for respiration streams through the clefts, gaseous interchange takes place between the circulating colourless blood and the percolating water. The pharynx projects freely into the atrium; it is surrounded at the sides and below by the continuous atrial cavity, but dorsally it is held in position in two ways. First, its dorsal wall (which is grooved to form the hyperpharyngeal groove) is closely adherent to the sheath of the notochord; and secondly, the pharynx is attached through the intermediation of the primary bars. These are suspended to the muscular bodywall by a double membrane, called the ligamentum denticulatum, which forms at once the roof of the atrial chamber and the floor of a persistent portion of the original body-cavity or coelom (the dorsal coelomic canal on each side of the pharynx). The ligamentum denticulatum is thus lined on one side by the epiblastic atrial epithelium, and on the other by mesoblastic coelomic epithelium. Now this ligament is inserted into the primary bars some distance below the upper limits of the gill-clefts, and it therefore follows that, corresponding with each tongue-bar, the atrial cavity is produced upward beyond the insertion of the ligament into a series of bags or pockets, which may be called the atrial pouches. At the top of each of these pouches there is a minute orifice, the aperture of a small tubule lying above each pouch in the dorsal coelom. These tubules are the excretory tubules or nephridia. They communicate with the coelom by several openings or nephrostomes, and with the atrium by a single opening in each case, the nephridiopore. It is important to emphasize the fact that in Amphioxus the excretory tubules are co-extensive with the gill-clefts. The perforated pharynx terminates some distance in front of the atriopore. At the level of its posterior end a pair of funnel-shaped pouches of the atrium are produced forwards into the dorsal coelom. These are the atrial coelomic funnels or brown funnels, so called on account of the characteristic pigmentation of their walls. There are reasons for supposing that these funnels are vestiges of an ancient excretory system, which has given way by substitution to the excretory tubules described above. In the same region of the body, namely, close behind the pharynx, a large diverticulum is given off from the ventral side of the gut. This is the hepatic caecum (fig. 2, 2, q, fig. 4, l&#8202;), which is quite median at its first origin, but, as it grows in length, comes to lie against the right wall of the pharynx. Although within the atrial cavity, it is separated from the latter by a narrow coelomic space, bounded towards the atrium by coelomic and atrial epithelium. No food passes into the hepatic caecum, which has been definitely shown on embryological and physiological grounds to be the simplest persistent form of the vertebrate liver.

Nervous System.—As has already been indicated, a solid sub-cylindrical elastic rod, the notochord, surrounded by a sheath of laminar connective tissue, the cordal sheath, lies above the alimentary canal in contact with its dorsal wall, and extends beyond it both in front and behind to the obtusely pointed extremities of the body. This notochord represents the persistent primordial skeletal axis which, in the higher Craniata (though not so in the lower), gives way by substitution to the segmented vertebral column. Immediately above the notochord there lies another