Experiments have proved the excellence of sugar as a substitute for honey, and in some instances its superiority for the formation of wax. It might otherwise have been supposed that bees might form comb from some particles of wax accidentally present in the honey, and that these afforded the pabulum for this secretion. To prove therefore that the saccharine principle alone enabled the bees to produce wax, being still confined, they were supplied with a syrup made with Canary-sugar and water, and at the same time comparative experiments were made in another hive, where the bees were fed on honey and water. The syrup-fed bees produced wax sooner and more abundantly than the honey-fed bees. Another fact was also incontrovertibly elicited; namely, that in the old hives the honey is warehoused, and that in the new ones it is consumed and transmuted into wax.
The experiments of Huber have been confirmed by those of M. Blondelu, of Noyau, who addressed a memoir upon this subject to the Society of Agriculture at Paris, in May 1812. Huish has critically examined these experiments of Huber, but without being convinced by them: for having observed pollen on the thighs of bees when swarming, and upon dissection, in their stomachs also, he considers that pollen, elaborated in the second stomach of the bee, “contains in itself the principle of wax.” Were this the case, what a store of pollen must the bees have reserved, in Huber’s experiments, wherein they formed five successive sets of comb, without access to fresh pollen! The pollen or bee-bread, which Huish discovered on the thighs and in the stomachs of some of his bees, was most likely intended for larva-food; they were probably bees that had been abroad, and joined the swarm on their passage home, before they had deposited their freight in the parent hive. With this pollen (or ambrosia, as it has been called), after conversion into a sort of whitish jelly by the action of the bee’s stomach, where it is probably mixed with honey, and then regurgitated, the young brood, immediately upon their exclusion and until their change into nymphs, are fed by the nursing-bees several times a day. The opinion that pollen is the prime constituent of wax was held by Buffon, and remains uncontradicted in an edition of his Works so late as 1821. Arthur Dobbs, Esq., in the Philosophical Transactions for 1752, instead of considering wax as digested pollen discharged from the stomach of the bee, regards it as being emitted per annum; and as he speaks of its discharge in husks or shells, doubtless he saw it in that form, which it is now known to assume when moulded upon the body of the bee. Indeed he says that he has had swarming bees alight upon his hand, and drop warm wax upon it. Its being secreted only by the under side of the belly might easily deceive, and lead him to regard it as alvine excrement.
I will here subjoin some more proofs of the non-identity of wax and pollen. So long ago as 1768, the Lusatian Society (called Société des Abeilles, founded at little Bautzen, a village in Upper Lusatia, under the auspices of the Elector of Saxony,) knew that wax was not discharged from the mouths of bees, but was secreted in thin scales among their abdominal rings or segments. About 1774, Mr. Thorley caught a bee just entering its hive, and found, among the plaits of its belly, no less than six pieces or scales of solid wax, perfectly white and transparent, and he oftentimes saw wax in the same situation. M. Duchet, in his Culture des Abeilles, quoted by Wildman in 1778, declares that wax is formed of honey; and relates in proof of it, that he has seen a broken comb of an overset hive, which was repaired during bad weather, when the bees could not acquire any other material. This statement of Duchet corresponds with my own observation, as stated in [page 357], but is not so conclusive. In Duchet’s instance there might have been other materials in the hive besides honey; whereas in my case the bees had access to no materials whatever, excepting the sugared ale and the honey which they had conveyed from the parent hive, the swarm having been just hived. Wildman, in his Treatise on the Management of Bees, states his having seen pieces of wax, like fish scales, on the hive floor of a fresh swarmed colony, part of which he thinks must at least have been formed upon the body of the bee; some flakes might have fallen from the combs then constructing, but there were many pieces among them which were concave on one side and convex on the other, as if moulded on the insect’s belly. Flakes were likewise seen, hanging loose, between the abdominal scales of the bees. In 1792, Mr. John Hunter, apparently unacquainted with antecedent conjectures, detected the genuine reservoir of wax under the bee’s belly. He considered wax as an external secretion of oil, formed and moulded between the abdominal scales of the insect. Dr. Evans confirms the testimony of Wildman and Hunter, having been an eye-witness to the formation of wax into flakes. “One or more bees,” he remarks, “may be often seen before the door of the hive, supporting themselves by their two fore-feet, fluttering their wings, and agitating the hind parts of their bodies. They are then evidently moulding the wax between their abdominal scales, the motion of the wings serving to preserve their balance, and as a signal for their companions within to come and carry off the falling flakes.” In the Philosophical Transactions for 1807, Mr. Knight states that there is no such secretory process; that the wax is laid on the scales of the abdomen for the convenience of carriage, and to receive warmth preparatory to cell-building.
To complete the evidence however, to me so irresistible, in favour of the wax-secreting faculty of the bee’s body, I observe finally, that in 1793, M. Huber’s observations led him to the same conclusion as Mr. Hunter’s, relative to the nature of the laminæ under the abdominal scales: but Huber slumbered not there, he prosecuted the inquiry more successfully than any preceding naturalist, and at length demonstrated the secreting organs which had eluded the scrutiny of Swammerdam, Hunter, and other acute anatomists. He found that these laminæ were contained in distinct receptacles, on each side of the middle process of the scales; he examined with great care the form and structure of these secreting cavities, which are peculiar to working bees. Each working bee has eight of these organs, sacklets or small compartments. Their general shape is an irregular pentagon, and the plates of wax, being moulded in them, exhibit accordingly the same form. A perforation of their lining membrane on the side next to the abdomen, started a jet of transparent fluid, which congealed on cooling; in this state it resembled wax, and became again fluid on the application of heat. Comparative experiments were made with the substance contained in the pouches and with the wax of fresh combs: a great similarity between these two substances was discerned; the latter appeared somewhat more compound, having probably received some additional ingredient, while employed as the material for building. The secreting function of the membrane on the inner surface of these cavities, was further evinced, by a more minute examination of its structure, which exhibited a number of folds, forming an hexagonal net-work, analogous to the inner coat of the second stomach of ruminating quadrupeds. Huber does not appear to have known the observations either of Duchet or of Wildman on this subject, although they were made long prior to Mr. Hunter’s; for he quotes only from the latter.
When combs are wanted, bees fill their crops with honey, and retaining it in them, hang together in a cluster from the top of the hive, and remain inactive about twenty-four hours. During this time the wax is secreted, and may be seen in laminæ, under the abdominal scales, whence it is removed by the hind legs of the bee, and transferred to the fore legs; from them it is taken by the jaws, and after being masticated as described in Chap, XXXIV, [page 347], the fabrication of comb commences.
“To see the wax-pockets in the hive-bee, you must press the abdomen, so as to cause its distention; you will then find, on each of the four intermediate ventral segments, separated by the carina or elevated central part, two trapeziform whitish pockets, of a soft membranaceous texture: on these the laminæ of wax are formed, in different states, more or less perceptible[AB].”
[AB] Kirby and Spence.
Messrs. Huber and Son ascertained that the office of collecting honey, for the elaboration of wax, is filled by a particular description of bees or labourers, to which they have given the name of wax-workers. These bees are susceptible of an increase in size, as is evident from the state of their stomachs, when quite full of honey. Dissection has shown that their stomachs are more capacious than those of the bees that are differently occupied. Bees not possessed of this expanding stomach, gather no more honey than is necessary to supply the immediate wants of themselves and their companions, with whom they readily share it: these are called nursing-bees, their principal duty being to attend the eggs and larvæ. The task of storing the hive with provisions devolves upon the wax-workers, who, when not occupied in the construction of comb, disgorge their honey into those cells which are intended for its reception. By marking the bees, it was found that they never encroached upon each other’s employment: this strict adjustment of duty is the more remarkable, since the power of producing wax is common both to the nursing- and wax-working bees, a small quantity of wax being really found in the receptacles of the nursing-bees.
In the foregoing experiments for ascertaining the sources of wax, the bees had borne their confinement without evincing the least impatience; but on another occasion, when shut up with a brood of eggs and larvæ, and without pollen, though honey was copiously supplied, they manifested uneasiness and rage at their imprisonment. Fearing the consequence of this state of tumult being prolonged, Huber allowed them to escape in the evening, when too late to collect provisions; the bees soon returned home. At the end of five days, during which this experiment was tried, the hive was examined:—the larvæ had perished, and the jelly that surrounded them on their introduction into the hive had disappeared. The same bees were then supplied with a fresh brood, together with some comb containing pollen: very different indeed was their behaviour with this outfit; they eagerly seized the pollen and conveyed it to the young; order and prosperity were re-established in the colony; the larvæ underwent the usual transformations; royal cells were completed and closed with wax, and the bees showed no desire to quit their habitation. These experiments afford indisputable evidence of the origin of wax and the destination of pollen.
Though the wax of honey and brood-comb be an original secretion from the body of the bee, wax is also considered by some as a vegetable substance existing abundantly in nature. According to Proust, it forms the silvery down on the leaves, flowers and fruit of many plants, and resides likewise in the feculæ of others. Dr. Darwin, in his Phytologia, supposes that wax is secreted to glaze over the fecundating dust of the anthers, and prevent its premature explosion from excessive moisture: to an unseasonable dispersion of anther-dust he ascribes the failure of orchard and corn crops in summers of extreme humidity. The wax-tree of Louisiana[AC] (Myrica cerifera) contains immense quantities of wax. In this respect there appears an identity betwixt animal and vegetable secretion, which may be viewed as indicative of simplicity in the structure of the bee: a still simpler organization exists in the aphis, which extracts the saccharine juices from the leaves and bark of trees, and expels them again nearly unchanged[AD].