Fig. 9. Larva of Ascidia mentula. (From Gegenbaur; after Kupffer.) Only the anterior part of the tail is represented.
N´. anterior swelling of neural tube; N. anterior swelling of spinal portion of neural tube; n. hinder part of neural tube; ch. notochord; K. branchial region of alimentary tract; d. œsophageal and gastric region of alimentary tract; O. eye; a. otolith; o. mouth; s. papilla for attachment.
The original single vesicle of the brain becomes divided by the time the larva is hatched into two sections ([fig. 9])—(1) an anterior vesicle with, for the most part, thin walls, in which unpaired auditory and optic organs make their appearance, and (2) a posterior nearly solid cephalic ganglion, through which there passes a narrow continuation of the central canal of the nervous system. This ganglion consists of a dorsal section formed of distinct cells, and a ventral section formed of a punctated material with nuclei. The auditory organ[8] consists of a ‘crista acustica’ ([fig. 9]), in the form of a slight prominence of columnar cells on the ventral side of the anterior cerebral vesicle; to the summit of which a spherical otolith is attached by fine hairs. In the crista is a cavity containing clear fluid. The dorsal half of the otolith is pigmented: the ventral half is without pigment. The crista is developed in situ, but the otolith is formed from a single cell on the dorsal side of the cerebral vesicle, which forms a projection into the cavity of the vesicle, and then travels (in a manner not clearly made out) round the right side of the vesicle till it comes to the crista; to which it is at first attached by a narrow pedicle. The fully developed eye ([figs. 8] VI. and [9], O) consists of a cup-shaped retina, which forms a prominence slightly on the right side of the posterior part of the dorsal wall of the anterior cerebral vesicle, and of refractive media. The retina is formed of columnar cells, the inner ends of which are imbedded in pigment. The refractive media of the eye are directed towards the cavity of the cerebral vesicle, and consist of a biconvex lens and a meniscus. Half the lens is imbedded in the cavity of the retina and surrounded by the pigment, and the other half is turned toward a concavo-convex meniscus which corresponds in position with the cornea. The development of the meniscus and lens is unknown, but the retina is formed ([fig. 8] V. a) as an outgrowth of the wall of the brain. At the inner ends of the cells of this outgrowth a deposit of pigment appears.
The trunk section of the spinal cord ([fig. 9], N) is separated by a sharp constriction from the brain. It is formed of a superficial layer of longitudinal nervous fibres, and a central core of ganglion cells. The layer of fibres diminishes in thickness towards the tail, and finally ceases to be visible. Kupffer detected three pairs of nerves passing off from the spinal cord to the muscles of the tail. The foremost of these arises at the boundary between the trunk and the tail, and the two others at regular intervals behind this point.
The mesoblast and muscular system. It has already been stated that the lateral walls of the archenteron in the tail give rise to muscular cells. These cells lie about three abreast, and appear not to increase in number; so that with the growth of the tail they grow enormously in length, and eventually become imperfectly striated. The mesoblast cells at the hinder end of the trunk, close to its junction with the tail, do not become converted into muscle cells, but give rise to blood corpuscles; and the axial remnant of the archenteron undergoes a similar fate. According to Kowalevsky the heart is formed during larval life as an elongated closed sack on the right side of the endostyle.
The notochord. The notochord was left as a rod formed of a single row of cells, or in As. canina and some other forms of two rows, extending from just within the border of the trunk to the end of the tail.
According to Kowalevsky, Kupffer, Giard, etc. the notochord undergoes a further development which finds its only complete parallel amongst Chordata in the doubtful case of Amphioxus.
There appear between the cells peculiar, highly refractive discs ([fig. 8] V. Chs). These become larger and larger, and finally, after pushing the remnants of the cells with their nuclei to the sides, coalesce together to form a continuous axis of hyaline substance. The remnants of the cells with their nuclei form a sheath round the hyaline axis ([fig. 8] VI. ch.). Whether the axis is to be regarded as formed of an intercellular substance, or of a differentiation of parts of the cells is still doubtful. Kupffer inclines to the latter view: the analogy of the notochord of higher types appears to me to tell in favour of the former one.
The alimentary tract. The anterior part of the primitive archenteron alone retains a lumen, and from this part the whole of the permanent alimentary tract (mesenteron) becomes developed. The anterior part of it grows upwards, and before hatching an involution of the epiblast on the dorsal side, just in front of the anterior extremity of the nervous system, meets and opens into this upgrowth, and gives rise to the permanent mouth ([fig. 8] V. o).
Kowalevsky states that a pore is formed at the front end of the nervous tube leading into the mouth ([fig. 8] V. and VI. f) which eventually gives rise to the ciliated sack, which lies in the adult at the junction between the mouth and the branchial sack. Kupffer however was unable to find this opening; but Kowalevsky’s observations are confirmed by those of Salensky on Salpa.
From the hinder end of the alimentary sack an outgrowth directed dorsalwards makes its appearance ([figs. 8] V. and [9], d), from which the œsophagus, stomach and intestine become developed. It at first ends blindly. The remainder of the primitive alimentary sack gives rise to the branchial sack of the adult. Just after the larva has become hatched, the outgrowth to form the stomach and œsophagus, etc. bends ventralwards and to the right, and then turns again in a dorsal and left direction till it comes close to the dorsal surface, somewhat to the left of and close to the hinder end of the trunk. The first ventral loop of this part gives rise to the œsophagus, which opens into the stomach; from this again the dorsally directed intestine passes off.