subcylindrical cord, also surrounded by a sheath of connective tissue. 
Fig. 5.—Diagram of embryo of Amphioxus seen from above in optical section. (Adapted from Hatschek.) pc, Praechordal head-cavity of embryo; cc, collar-cavity (first somite); my, mesodermic somites (myocoelomic or archenteric pouches); ch, notochord with the neural tube (neurochord) lying upon it; np, anterior neuropore; ne, position of posterior neurenteric canal.This cord is neither elastic nor solid, but consists of nerve tissue, fibres and ganglion cells, surrounding a small central canal. For the sake of uniformity in nomenclature this nerve-cord may be called the neurochord. It is the central nervous system, and contains within itself the elements of the brain and spinal marrow of higher forms. The neurochord tapers towards its posterior end, where it is coextensive with the notochord, but ends abruptly in front, some distance behind the tip of the snout. The neurochord attains its greatest thickness not at its anterior end but some way behind this region; but the central canal dilates at the anterior extremity to form a thin-walled cerebral vesicle, in the front wall of which there is an aggregation of dark pigment cells constituting an eyespot, visible through the transparent skin (fig. 1). There are two pairs of specialized cerebral nerves innervating the praeoral lobe, and provided with peripheral ganglia placed near the termination of the smaller branches. Corresponding with each pair of myotomes, and subject to the same alternation, two pairs of spinal nerves arise from the neurochord, namely, a right and left pair of compact dorsal sensory roots without ganglionic enlargement, and a right and left pair of ventral motor roots composed of loose fibres issuing separately from the neurochord and passing directly to their termination on the muscle-plates of the myotomes. The first dorsal spinal nerve coincides in position with the myocomma which separates the first myotome from the second on each side, and thereafter the successive dorsal roots pass through the substance of the myocommata on their way to the skin; they are therefore septal or intersegmental in position. The ventral roots, on the contrary, are myal or segmental in position. In addition to the cerebral eyespot there are large numbers of minute black pigmented bodies beside and below the central canal of the neurochord, commencing from the level of the third myotome. It has been determined that these bodies are of the nature of eyes (Becheraugen, R. Hesse), each consisting of two cells, a cup-shaped pigment cell and a triangular retinal cell. These may be called the spinal eyes, and it is said that they are disposed in such a way as to receive illumination preferentially from the right side, although this fact has no relation with the side upon which Amphioxus may lie upon the sand. When kept in captivity the animal often lies upon one side on the surface of the sand, but on either side indifferently. Over the cerebral eye there is a small orifice placed to the left of the base of the cephalic fin, leading into a pit which extends from the surface of the body to the surface of the cerebral vesicle; this is known as A. von Kölliker’s olfactory pit.
Reproductive System.—The sexes are separate, and the male or female gonads, which are exactly similar in outward appearance, occur as a series of gonadic pouches projecting into the atrial cavity at the base of the myotomes (figs. 2, 3, 4). At the breeding season the walls of the pouches burst and the sexual elements pass into the atrium, whence they are discharged through the atriopore into the water, where fertilization takes place.
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| Fig. 6.—Anterior region of two pelagic larvae of A. lanceolatus obtained by the tow-net in 8–10 fathoms, showing the asymmetry of the large lateral sinistral mouth with its ciliated margin cm and the dextral series of simple primary gill-slits (1ps-14ps). The larvae swim normally like the adult or suspend themselves by their flagella (not shown in the figures) vertically in mid-water. There is nothing in their mode of life which will afford an explanation of the asymmetry which is a developmental phenomenon. Lettering of upper figure.—anp, Anterior neural pore; bc, rudiment of buccal skeleton; c, cilia; cb, ciliated band; cc, ciliated groove; cm, cilia at margin of mouth; gl, external opening of club-shaped gland; Hn, Hatschek’s nephridium; lm, left metapleur; n, notochord; pp, praeoral pit; ps, primary gill-slits, 1, 5, and 13; rm, right metapleur showing through. Lettering of lower figure.—a, Atrium; al, alimentary canal; bv, blood-vessel; cv, cerebral vesicle; df, dorsal section of myocoel (=fin spaces); e, “eyespot”; end, endostyle; gl, club-shaped gland; lm, edge of left metapleur; m, lower edge of mouth; n, notochord; nt, pigmented nerve tube; ps, primary gill-slits, 1, 9, and 14; rc, renal cells on atrial floor; rm, edge of right metapleur; so, sense organ opening into praeoral pit; ss, thickenings, the rudiments of the row of secondary gill-slits. |
Development.—The development of Amphioxus possesses many features of interest, and cannot fail to retain its importance as an introduction to the study of embryology. The four principal phases in the development are: (1) Blastula, (2) Gastrula, (3) Flagellate Embryo, (4) Larva. The segmentation or cleavage of the ovum which follows upon fertilization terminates in the achievement of the blastula form, a minute sphere of cells surrounding a central cavity. Then follows the phenomenon of gastrulation, by which one-half of the blastula is invaginated into the other, so as to obliterate the segmentation cavity. The embryo now consists of two layers of cells, epiblast and hypoblast, surrounding a cavity, the archenteron, which opens to the exterior by the orifice of invagination or blastopore. One important fact should be noted with regard to the gastrula, in which it seems to differ from the gastrulae of invertebrata. After invagination is completed, the embryo begins to elongate, the blastopore becomes narrower, and the dorsal wall of the gastrula loses its convexity, and becomes flattened to form the dorsal plate, the outer layer of which is the primordium of the neurochord and the inner layer the primordium of the notochord. While still within the egg-membrane the epiblastic cells become flagellated, and the gastrula rotates within the membrane. About the eighth hour after commencement of development the membrane ruptures and the oval embryo escapes, swimming by means of its flagella at the surface of the sea for another twenty-four hours, during which the principal organs are laid down, although the mouth does not open until the close of this period. The primordium of the neurochord (neural or medullary plate) referred to above becomes closed in from the surface by the overgrowth of surrounding epiblast, and its edges also bend up, meet, and finally fuse to form a tube, the medullary or neural tube. An important fact to note is that the blastopore is included in this overgrowth of epiblast, so that the neural tube remains for some time in open communication with the archenteron by means of a posterior neurenteric canal. It is still longer before the neural tube completes its closure in front, exhibiting a small orifice at the surface, the anterior neuropore. It is thus possible that the neurenteric canal is due to the conjunction of a posterior neuropore with the blastopore, i.e. it is a complex and not a simple structure. Paired archenteric pouches
