LITERATURE ON THE SALIVARY GLANDS

Leydig, F. Zur Anatomie der Insekten. (Archiv Anat. und Phys. 1859.)

—— Untersuchungen zur Anatomie und Histiologie der Tiere. Bonn, 1883, pp. 174, 8 Taf.

—— Intra- und interzellulare Gänge. (Biolog. Centralblatt, x, 1890, pp. 392–396.)

Dohrn, A. Zur Anatomie der Hemipteren. (Stettin. Entom. Zeit., 1866, salivary glands, pp. 328–332.)

Kupffer, C. Die Speicheldrüsen von Periplaneta orientalis und ihr Nervenapparat. (Beiträge zur Anatomie und Physiol., 1875.)

Schiemenz, P. Ueber das Herkommen des Futtersaftes und die Speicheldrüsen der Biene. (Zeitschr. f. wissens. Zool., xxxviii, 1883, pp. 71–135, 3 Taf.)

Korschelt, E. Ueber die eigentümlichen Bildungen in den Zellkernen der Speicheldrüsen von Chironomus plumosus. (Zool. Anzeiger, 1884, pp. 189–194, 221–225, 241–246.)

Hofer, B. Untersuchungen über den Bau der Speicheldrüsen und des dazu gehörenden Nervenapparates von Blatta. (Nova Acta d. Kais. Leopold.-Carol. Deutsch. Akad. d. Naturforscher, li, 1887, pp. 345–395, 3 Taf.)

Knüppel, A. Ueber Speicheldrüsen von Insekten. (Archiv für Naturg., 1887, Jahrg. 52, pp. 269–303, 2 Taf.)

Blanc, Louis. La tête du Bombyx mori à l’état larvaire, anatomie et physiologie. (Extrait des Travaux du Laboratoire d’Études de la Soie, 1889–1890; Lyon, 1891, p. 180, many figs.)

Bordas, L. Anatomie des glandes salivaires des Hyménoptères de la famille des Ichneumonidæ. (Zool. Anzeiger, 1894, pp. 131–133.)

—— Glandes salivaires des Apides, Apis mellifica. ♂ and ♀. (Comptes rendus Acad. Sc., Paris, cxix, pp. 363, 483, 693–695, 1894; also two articles in Bull. Soc. Philomath. Paris, 1894, pp. 5, 12, 66.)

—— Appareil glandulaire des Hyménoptères. (Ann. Sc. Nat. Zool., xix, Paris, 1894, pp. 1–362, 11 Pls.) (See also p. 366.)

Berlese, Antonio. Le cocciniglie Italiane viventi sugli agrumi. Firenze, 1896, 12 Pls. and 200 Figs.

With the writings of Mark, Minot, Locy, List, Krassilstschik, Nagel (1896).

b. The silk or spinning glands, and the spinning apparatus

The larvæ of certain insects, chiefly those of the Lepidoptera, possess a pair of silk or spinning glands (sericteries) which unite to form a single duct opening in the upper lip at the end of the lingua, which is modified to form the spinneret. (See pp. 71, 75.) All caterpillars possess them, and they are best developed in the silkworms, which spin the most complete cocoon. Silk-glands also occur in the larvæ of the Tenthredinidæ, in the case-worms or larval Trichoptera, also in certain chrysomelid beetles (Donacia, Hæmonia), and in a weevil (Hypera). In a common caddis-worm (Limnophilus) the glands are of a beautiful pale violet-blue tint, and two and a half times as long as the larva itself; viz. the body is 20 mm. and the glands 55 mm. in length.

In caterpillars the glands are of tubular shape, shining white, and much like the ordinary simple tubular salivary glands of the imago. When only slightly longer than the body they are twice folded, the folds parallel and situated partly beneath and partly on the side of the digestive canal; not usually, when folded in their natural position, extending much behind the end of the stomach; but in the silkworms they are so long and folded as to envelop the hinder part of the canal. In geometrid caterpillars the glands when stretched out only reach slightly beyond the end of the body; in Datana they are half again as long as the body. Helm thus gives their relative length in certain Eurasian caterpillars, and we add that of Telea polyphemus:—

Vanessa iolength of body32 mm.; of the silk glands26 mm.
Smerinthus tiliælength of body63 mm.; of the silk glands205 mm.
Bombyx morilength of body56 mm.; of the silk glands262 mm.
Antheræa yamamayalength of body100 mm.; of the silk glands625 mm.
Telea polyphemuslength of body60 mm.; of the silk glands450 mm.

Thus in Telea the silk-glands are about 18.50 inches in length, being about seven times as long as the body.

For the most complete accounts of the spinning glands of Lepidoptera and their mechanism we are indebted to Helm and to Blanc, and for that of the Trichoptera to Gilson.

The unpaired portion, or spinning apparatus (filière of Lyonet), is divided into two portions; the hinder half being the “thread-press,” the anterior division the “directing tubes.” The silk material, stored up in the thickest portion of the glands, passes into the thread-press (Fig. 334, A), which is provided with muscles which force the two double ribbon-like threads through the directing tube, as wire is made by molten iron being driven through an iron plate perforated with fine holes. The entire spinning apparatus, or filator, as we may call it, is situated in the tubular spinneret. The opening of the spinneret is directed anteriorly, and the anterior end of the directing tube passes directly into this opening so that the directing tube may be regarded as an invagination of the lingua.

The silk thread which issues from the mouth of the spinneret is, as Leeuwenhoek discovered, a double ribbon-like band, as may be seen in examining the silk of any cocoon.

The process of spinning.—Since the appearance of Helm’s account, Gilson, and also Blanc, have added to our knowledge of the way in which the silk is spun and of the mechanism of the process. Gilson has arrived, in regard to the function of the press or filator, at the following conclusions: 1, the press regulates the thread, it compresses it, gives it its flattened shape; 2, it regulates the layer of gum[[52]] (grès) which surrounds the thread; 3, it may render the thread immovable by compressing it as if held by pincers.

The process of spinning in the silkworm, says Blanc, comprises all the phenomena by which the mass of silk contained in the reservoir is transformed into the silk fluid of which the cocoon is spun. The excretory canals each contain a cylindrical thread of silk having a mean diameter of 0.2 mm. and surrounded by a layer of gum (grès) which in the fresh living organ exactly fills the annular space situated between the fibroin cylinder and the wall. Arrived within the common duct, the two threads receive the secretion of Filippi’s gland, where the silken fluid is formed, but has not yet assumed its definite external characters. The two threads press through the common canal and arrive at the infundibulum (Fig. 334, c) of the press, at the bottom of which is situated the orifice of the spinning canal, almost completely divided into two by the sharp edge of the rachis (Figs. 334, a, 335, l). The threads each pass into one of the two grooves, and the layer of gum (grès) fills the rest of the canal of the press or filator.

Fig. 331.—Longitudinal section of the spinneret: a, horizontal portion of the tongue; b, vertical portion; c, f, circle of the tongue; d, tongue-pad; e, orifice of the spinneret; g, body of the lyre; h, prebasilar membrane forming a fold; i, internal canal of the spinneret; k, filator.—After Blanc.

Fig. 332.—The lower lip (labium) of Bombyx mori, isolated, seen from the left side: A, lyre; B, spinneret; C, labial palpus; D, vertical part of the labium; E, horizontal part of the same; H, L, silk-canal; K, right gland of Filippi; L, canal of the left gland; N, labial nerve; a, oblique fibre of the elevator of the labium; b, right fibre of the same; c, depressor of the labium; d, superior spinning muscles.

Fig. 333.—The labium in a horizontal position, seen from the side: f, the filator or press situated under the external part of the spinneret (d), between the branches (b), of the lyre (a); e, labial palp; c, tongue.

Fig. 334.—Longitudinal section of the spinneret and press (filator): A, filator or press; B, spinneret; C–D, body of the lyre; F, lower part of the labium; E, common canal; eh, its epithelium; G, superior muscle of the press; a, rachis; b, its posterior enlargement; c, infundibulum; d, cuticle; o, orifice of the spinning canal; op, central canal of the lyre and of the spinneret; fi, hypodermis of the lyre; f, f, hypodermic pad of the lyre.

The silken substance is then pressed by the more or less powerful contractions of the muscles of the filator, so that the passage of the threads is facilitated. If the muscles totally contract, the spinning canal is opened wide, the threads pass easily upwards and assume the form of a triangular prism (Fig. 336).

Fig. 335.—Spinning apparatus, seen from above: A, opening of the spinneret; B, central canal of the spinneret (C); D, common canal; E, canal of Filippi; F, excretory canal of a silk-gland; i, orifice of the canal of Filippi’s gland; l, rachis; k, ring of the infundibulum; b, c, d, e, f, cavity of the different canals; h, spur which separates the two excretory canals.—This and Figs. 331–334 after Blanc.

If this contraction diminishes, the chitinous wall of the spinneret comes together, owing to its elasticity; the ceiling of the canal approaches the floor; the cavity tends to take the form of a semicircular slit, and the threads are compressed, flattened. As each mass or thread of silk is much more voluminous than the canal, except when the latter is extremely dilated, it follows that the two threads are always compressed, or squeezed together, and that each of them is compelled to mould itself in the groove it occupies and to take its shape. Hence the variations in the appearance of the two masses or divided portions of silk, which as stated present all grades between the form of an isosceles-triangular prism and that of a nearly flat ribbon; but this last case is quite rare. The use of the spinneret, then, is to compress the thread and to change its form more or less considerably, at the same time as it diminishes its diameter.

Fig. 336.—Diagram of the press and its muscles: a, lower; b, lateral; c, upper muscles of the press.—After Blanc.

Moreover, this constant compression of the thread as it passes through the press keeps it in a certain state of tension so as to allow the caterpillar while spinning to firmly hold its thread.

Finally, when the worm suspends the contraction of its spinning muscles, the press flattens, vigorously compresses the thread, and arrests its motion, in such a way that if there was a strain on the silken fluid (bave), it would break rather than oblige the caterpillar to let go any more of it.

The press does not act directly on the silken thread, but through the gummy layer (grès) which transmits over the whole surface of the silken fluid (brin) the pressure exerted on it. After having overcome this difficult passage, the silk thread has acquired its definite form; it rapidly passes out of the spinneret.

How the thread is drawn out.—Having seen, says Blanc, how the two masses of silk (brins), in passing through the spinning apparatus (or press), join each other, constituting the frothy silken fluid, thus becoming modified in form, it remains to examine the way in which the thread is drawn out of the spinneret. If we examine a caterpillar while spinning, it will be seen that in moving its head it draws on the frothy mass of silk fixed to the web of the cocoon. This traction certainly aids very much the exit of the thread, but it is not the only cause.

The silk, Blanc affirms, is pushed out by a force a tergo, developed by different agents, such as the pressure of the distended cuticle or the silky mass contained in the reservoir, as seen in the section of a worm which has spun its cocoon. But if we consider a caterpillar before it has begun to spin, it is difficult to explain the mechanism of spinning. As Blanc has often observed, in making sections of the heads of silkworms, two cases arise. Sometimes the worm has already spun a little, and a certain length of the frothy silk (bave) issues from the orifice of the spinneret, where it forms a small twisted bundle. At other times the worm has not spun since its last moult or the frothy mass of silk has broken within the head, and we find the end in the common tube. In the first of these two cases, the worm, dilating its press, is able by a general contraction to discharge a little of the gritty material (grès) which lines the ball of silk hanging at the end of the spinneret. It can also reject a certain quantity of the secretion of Filippi’s glands and thus soften the gritty substance. The little plug of silk can then adhere to the body with which it comes in contact.

In the same case it is necessary that the two bits or portions of silk traverse the press, and this normally has a calibre less than their diameter. The worm should then distend the spinning tube as much as is practicable, so as to make the openings as large as possible. It has been stated that the press is, in this condition, at least as large as the mass of frothy silk. This Blanc believes (although Gilson thinks otherwise) is pushed by a force a tergo, and reaches the funnel of the spinning canal; its two bits of silk (brins) unite there, penetrate into the canal itself, and, owing to successive impulses produced by the general contractions of the worm, press through and pass out of the spinneret.

While the silkworm is engaged in spinning its cocoon, the spinneret and press execute very varied movements, determined by the elevator, depressor, retractor, and protractor muscles of the labium, as well as those of the press. These movements, originally very numerous, may combine among themselves, so that the spinneret is susceptible of assuming during the process of spinning still more diverse positions.

Fig. 337.—Portion of the silk-gland of Bombyx mori: p, tunica propria; i, tunica intima; s, secretion-cell with branched nuclei; a, separate secretion-cell from the anterior part of the silk-gland of Amphidasis betularia; b, the same of Vanessa urticæ; c, the same in Smerinthus tiliæ.—After Helm.

Fig. 338.—A, section of gland of lepidopter: B, section of silk-gland of a saw-fly larva; n, nucleus; i.d, canals; d.s, cavity.—After Gilson.

Histologically the silk-glands are composed of three layers,—the outer, or tunica propria (Fig 337); the inner, the tunica intima; the middle layer being composed of extraordinarily large epithelial cells which can be seen with the naked eye, and are also remarkable for the branched shape of the nuclei (a, b, c, 337), the branches being more or less lobed, and the larger the cells the more numerous are the branches of the nucleus. Gilson[[53]] finds that those of Trichoptera, Lepidoptera, Diptera, and Hymenoptera ordinarily consist of a small number of cells; and it is quite common, he says, to find only two cells in a transverse section (Fig. 338, A). In the Tenthredinidæ, however, “the organ still consists of a tube, the wall of which is composed of flat cells, but in addition to that, two series of spheroidal cells are attached to the sides. Each of these cells contains a system of tiny canals running through their cytoplasm (B, i. d). These cells are the secreting elements; they continually cast the silk substance into the tube.” A peculiarity of the tunica intima is its distinct transverse striation.

Fig. 339.—Branching nucleus of spinning gland of Pieris larva.—After Korschelt, from Wilson.

Fig. 340.—Filippi’s glands (G) isolated and seen from above: e, e, its lobules; d, its excretory canal; E, silk-duct; C, common canal; c, upper spinning muscle; b, lower muscle; a, lateral muscle; T, spinneret.—After Blanc.

The lining of the glands and of their common duct is moulted when the caterpillar casts its skin, and this, as well as the mode of development, shows that the glands are invaginations of the ectoderm. Gilson finds that the silk-glands and silk-apparatus of Trichoptera are very similar to those of caterpillars, and that the silk is formed in the same way.

Appendages of the silk-gland (Filippi’s glands).—In most larvæ there is either a single or a pair of secondary glands which open into the spinning glands near their anterior end. They are outgrowths of the gland provided with peculiarly modified excretory cells or evaginations of the entire glandular epithelium. Those of Bombyx mori (Fig. 340) are very well developed, and, according to Blanc, form two whitish, lobulated masses in the labium on each side of the common duct of the spinning gland. Externally they appear to be acinose; but their structure, as described by Blanc and by Gilson, is very peculiar. Helm thinks, with Cornalia, that the function of these glands is to secrete the adhesive fluid which unites the silk threads, and also to make the silk more adhesive in the process of spinning, but Blanc states that this is done before the thread passes into the common excretory tubes, and he is inclined to think that the secretion serves to lubricate the spinneret, and thus to facilitate the passage of the thread. On the other hand, in certain caterpillars these glands are situated quite far from the spinning apparatus.

The silk-glands in the pupa state undergo a process of degeneration, and finally completely disappear. They are specific larval organs evolved in adaptation to the necessity of the insect’s being protected during its pupal life by a cocoon. (Helm.)

Morphologically the silk-glands are by Lang regarded as modified coxal glands, and homologues of the setiparous parapodial glands of chætopod worms, the coxal glands of Peripatus, and the spinning glands of spiders.

In Scolopendrella, spinning glands are situated in the two last segments of the body, opening out at the end of the cercopods (Fig. 15, s.gl), and the larvæ of the true Neuroptera (Chrysopa, Myrmeleon, etc.) which spin cocoons, have spinning glands opening into the rectum. The silk forming the cocoon of the ant-lion, as Siebold and the older observers have stated, is secreted by the walls of the rectal or anal sac. Siebold (Anatomy of the Invertebrates, p. 445) states that in the larva of Myrmeleon, the silk-apparatus is very remarkable, “for the rectum itself is changed into a large sac and secretes this substance which escapes through an articulated spinneret projecting from the opening of the anus”[[54]] (Fig. 307, e). The larvæ of the Mycetophilidæ have spinning glands at the hinder end of the body, as also the imago of the female of the tineid moth Euplocamus. (Kennel.) The larvæ of ichneumons, wasps, bees, of Cecidomyia, and other Diptera, spin silken cocoons, but their glands have not yet been examined.

It should also be observed that during the process of pupation the larvæ of butterflies, of certain flies (Syrphus), and beetles (Coccinellidæ and some Chrysomelidæ) attach themselves by silk spun from the anus, so that the pupa is suspended by its tail; such glands are probably homogenetic with the coxal glands.

The silk in its fluid or soft state is mucilaginous, and according to Mulder, in the silkworm consists of the following substances, varying somewhat in their relative proportions by weight:

Silk-fibre material53.67
Glue (Leim)20.66
Protoplasm24.43
Wax1.39
Coloring matter0.05
Fat and resin0.10