Structure.—The more important points in the structure of a typical Salpa are shown in Fig. 65. The branchial and atrial apertures are at opposite ends of the body, and lead into large cavities, the branchial and peribranchial sac respectively, which are in free communication at the sides of the obliquely-running dorsal lamina or "gill" (d.l). The transparent test is usually thick, and varies from a gelatinous to a stiff cartilaginous condition; it adheres closely to the surface of the mantle (ectoderm and body-wall). The muscle-bands (from 4 to about 20—usually 8 or 10) of the mantle do not in most cases completely encircle the body. They are present dorsally (Fig. 65, mus.bds) and laterally, but the majority do not reach the ventral surface. In many cases neighbouring bands join in the median dorsal line (Fig. 61). The muscle fibres are striated, and have rows of large equidistant nuclei. The anterior end of the dorsal lamina is in some cases prolonged to form a prominent tentacular organ, the languet or dorsal tentacle, projecting into the branchial sac, while near this opens a ciliated funnel corresponding to the dorsal tubercle, but having no connexion in the adult with either ganglion or subneural gland. The conjoined ganglion and subneural gland, the dorsal lamina, the peripharyngeal bands and the endostyle are placed in the usual positions. Eyes in the form either of a continuous horse-shoe-shaped pigmented ridge on the dorsal surface of the ganglion immediately below the ectoderm, or of one larger median and several smaller lateral ocelli are found in the various species of Salpa. These eyes have in most cases a retina formed of elongated cells, and a pigment-layer placed upon the ganglion.
The so-called otocysts of Salpa have been shown by Metcalf to be really glandular organs. They have been called lateral neural glands; they do not open at the dorsal tubercle, but separately into the pharynx. These lateral neural tubular glands have also been regarded as nephridia.
The large spaces at the sides of the dorsal lamina (often called the gill or branchia of Salpa), by means of which the cavity of the branchial sac is placed in free communication with the peribranchial cavity, are to be regarded as gigantic gill-slits formed by the suppression of the lateral walls and small stigmata of the branchial sac. The alimentary canal at the posterior end of the "gill" consists of oesophagus, stomach, and intestine, with a pair of lateral gastric glands or caeca. These viscera along with the reproductive organs, when present, make up the "nucleus" (Fig. 66, v).
Fig. 65.—Diagrammatic sagittal section of a "chain" Salpa. an, Anus; at, atrial aperture; at.m, muscles of atrial aperture; atr.cav, atrial cavity; br, branchial aperture; br.m, muscles of branchial aperture; br.s, branchial sac; d.l, dorsal lamina or "gill"; d.t, dorsal tubercle; end, endostyle; ht, heart; int, intestine; l, sensory languet; mus.bds, muscle-bands; n.g, nerve-ganglion; oc, eye-spot; oe, oesophagus; ov, ovary; p.p.b, peripharyngeal band; s.gl, neural gland; stom, stomach; t, t′, test; tes, testis; z, prebranchial zone. (After Herdman.)
Alternation of Generations.—Fig. 66 represents an aggregated or sexual Salpa, which was once a member of a chain, since it shows a testis and a developing embryo. The ova (always few in number, usually only one) appear at a very early period in the developing chain Salpa, while it is still a part of the gemmiparous stolon in the body of the solitary Salpa. This gave rise to the view put forward first by Brooks that the ovary really belongs to the solitary stolon-bearing Salpa, which is therefore a female producing a series of males by asexual gemmation, and depositing in each of these an ovum, which will afterwards, when fertilised, develop in the body of the male into a solitary or female Salpa. This idea, if adopted, would profoundly modify our conception of Salpa as an example of a life-history showing alternation of generations, but it seems to me to give a distorted view of the sequence of events. The fact that the stolon while in the solitary Salpa contains, along with representatives of other important systems of the body, a row of germinal cells, does not constitute that solitary Salpa the parent of the ova which these germinal cells will afterwards become in the body of an independent bud. We must regard as the parent the body in which the ova become mature and fulfil their function. The sexual or chain Salpa, although really hermaphrodite in its life-history, is usually[[109]] protogynous, i.e. the ova mature at an earlier period than the male organ or testis. This prevents self-fertilisation. The ovum is presumably fertilised by the spermatozoa of an older Salpa belonging to another chain, and the embryo is far advanced in its development before the testis is formed. The development takes place inside the body of the parent, and is "direct"—no tailed larval form being produced.
Fig. 66.—Salpa hexagona, Q. and G. Chain form dissected from the left side. a, Anus; at, atrial aperture; br, branchial aperture; d.l, dorsal lamina ("gill"); d.t, dorsal tubercle; emb, embryos; end, endostyle; m.b 2, m.b 7, second and seventh muscle-bands; n.g, nerve-ganglion; v, visceral "nucleus." (After Traustedt.)
Development and Life-history.—The segmentation of the egg is holoblastic, and gives rise to a number of blastomeres, which are for a time masked by the phenomenal activity of certain cells of extraneous origin, the "kalymmocytes," derived from the follicular epithelium surrounding the ovum. These follicular kalymmocytes migrate into the ovum, surround groups of blastomeres, and arrange themselves so as to reproduce the essential structure of the future embryo for which they form what may be termed a scaffolding or temporary support. After a time the blastomeres become active, proliferate rapidly, and finally press upon and absorb the kalymmocytes, and so eventually take their proper place in building up the organs. Some observers regard the kalymmocytes as being passive and nutritive only in function.
