Origin—Production—Chemical Constitution—Comb Building—Detailed Description—Amount of Wax in Hives—Commercial Value—Properties.
It was long thought that wax was a product derived, like honey, immediately from flowers. Not only did popular ignorance suppose that the pellets of pollen carried on the thighs of the worker-bees consisted of this substance, but even some authors on apiculture fell into the same error. It is now ascertained with certainty that wax is a sort of animal fat, elaborated from honey by certain internal organs of the bee. It exudes in a liquid form from sacklets on the under side of each of the four intermediate ventral segments of the abdomen. There are two of these pockets to each segment, one on either side of the carina or elevated central part. They are trapeziform in shape, and impart the same form to the tiny plates which emerge from them. On reaching the air the liquid thickens, and dries in flakes like fish-scales. The secretion of wax is carried on by the workers only, queens and drones being destitute of the apparatus necessary for the purpose. No direct communication has been traced between the stomach and the wax-sacks, but it has been conjectured by Hunter that the secretion is effected by the network of vessels lining the receptacles as a membrane covered with hexagonal cells, somewhat like the second stomach of ruminating quadrupeds.
Chemically considered, wax consists entirely of carbon, oxygen, and hydrogen; and, as before mentioned, is elaborated wholly from honey. Some authors have maintained that pollen is necessary for its production, but this is the case probably only indirectly; that is to say, the nitrogenous constituent of pollen may be necessary for the nutriment and stimulation of the secreting organs. It certainly does not enter into the constitution of the wax itself.
The quantity of honey required for this process of wax-making is very large. It is generally believed, in fact, to be from fifteen to twenty times the weight of the material derived from it; in other words, for every ounce of wax produced, at least a pound of honey is consumed by the bees. During the oxygenation of so large a quantity of saccharine matter, much heat is evolved a fact frequently noticed when comb-building is going on rapidly in a hive.
When wax is required for the abode of a fresh swarm, or for filling up vacant spaces with comb, the bees hang in festoons or chains, crossing the hive in different directions. Remaining almost motionless for about twenty-four hours, the wax-makers proceed with their business. Then, as soon as the little scales are of the proper consistency, they are withdrawn by the pincers on the hind legs of the bee, and carried between the fore-legs to the mouth. There, worked up with a small quantity of saliva, the substance is softened ready for use, and being conveyed away by those who have prepared it, and deposited in small masses, it furnishes the materials from which the comb-builders do their share of the duties of the hive. Possibly some of the individuals of the lower parts of the festoons, or clusters, may pass up their portions of wax to those above them for transmission to the top of the hive; but the fact is not thoroughly ascertained. Evans graphically says:—
It often happens that the fine scales fall by accident, or perhaps, when superabundant in quantity, on to the floor-boards of hives, and it does not appear, from our observation, that those bees who happen to come upon these little portions of material carry them up for employment in cell-formation.
The wax is used in comb-building, and the subject is one of great interest on many accounts, but especially from the following considerations: the nature of the material; the organs by which it is produced; the implements with which it is fashioned into shape; the manner in which the work is done; the form of the cells, the mathematical characters of which are most surprising; their different sizes and shapes, according to the purposes for which they are destined; their perfect adaptation to the needs of the bee community.
With regard to the nature of the material, in addition to the facts already mentioned, we may note that it is a substance easily moulded, especially when exposed to a gentle heat, such as is generated in a hive. It is light, so as to add little to the weight of the contents which will be stored in the cells. It is also a very slow conductor of heat, a matter of great importance both in summer and in winter. For, if it readily both absorbed and radiated heat, the temperature would, in the former season, become too high; while, in the winter, too great effort, and a large additional amount of food, would be needed by the bees to keep up the temperature of the hive to a point of safety for its inhabitants. Again, wax is a material which, by means of propolis (of which we shall presently speak), admits of being fastened in position so securely as to be able to bear a great weight of brood, honey, and bee-bread, in the cells.
The organs by which wax is secreted, and the implements with which it is fashioned, will be described fully in the chapter devoted to the physiology and anatomy of the bee; but we may say here that they are exceedingly simple, and that it is wonderful such beautiful work can be accomplished by means of them.
But the manner in which comb-building is done is so marvellous, that it merits a detailed description. It is to Huber that we are indebted for the full exposition of this subject, and we cannot do better than quote his account of the process, as given by Kirby and Spence. We must premise, however, that the great naturalist thought there were two distinct classes of workers, the one of which he called the wax-makers; the other, the nurse-bees. Observations continued since his day have rendered it certain that this is a mistaken distinction. As a general rule the care of the young devolves, as we have already said, on the most recently hatched of the community, who are unfit, for some days after emerging from the cell, to take distant flight in search of stores from flowers. The older and stronger workers, on the other hand, go abroad for supplies, and then, on their return, secrete whatever wax is needed in the economy of the hive.
The process of comb-building is described by Huber as follows:—"The wax-makers having taken a due portion of honey or sugar, from either of which wax can be elaborated, suspend themselves to each other, the claws of the fore-legs of the lowermost being attached to those of the hind pair of the uppermost, and form themselves into a cluster, the exterior layer of which looks like a kind of curtain. This cluster consists of a series of festoons or garlands, which cross each other in all directions, and in which most of the bees turn their back upon the observer.... The wax-makers remain immovable for about twenty-four hours, during which period the formation of wax takes place, and thin laminæ of this material may be generally perceived under their abdomen.
"One of these bees is now seen to detach itself from one of the central garlands of the cluster, to make a way amongst its companions to the middle of the vault, or top of the hive, and by turning itself round to form a kind of void, in which it can move itself freely. It then suspends itself to the centre of the space which it has cleared, the diameter of which is about an inch. It next seizes one of the laminæ of wax with a pincer formed by the posterior metatarsus (last joint of the leg), and tibia (last joint but two), and drawing it from beneath the abdominal segments, one of the anterior legs takes it with its claws and carries it to the mouth. This leg holds the lamina with its claws vertically, the tongue rolled up serving for a support, and by elevating it or depressing it at will, causes the whole of its circumference to be exposed to the action of its mandibles (or jaws), so that the margin is soon gnawed into pieces, which drop as they are detached into the double cavity, bordered with hairs, of the mandibles (jaws). These fragments, pressed by others newly separated, fall on one side of the mouth, and issue from it in the form of a narrow riband. They are then presented to the tongue, which impregnates them with a frothy liquor like a bouilli. During this operation the tongue assumes all sorts of forms; sometimes it is flattened like a spatula, then like a trowel, which applies itself to the riband of wax. At other times it resembles a pencil terminating in a point. After having moistened the whole of the riband, the tongue pushes it to make it re-enter the mandibles, but in an opposite direction, where it is worked up anew. The liquor mixed with the wax communicates to it a whiteness and opacity which it had not before; and the object of this mixture, which did not escape the observation of Réaumur, is, doubtless, to give it that ductility and tenacity which it possesses in its perfect state.
"The foundress-bee—the name which this first beginner of a comb deserves—next applies these prepared parcels of wax against the vault (or top of a frame) of the hive, disposing them with the point of her mandibles in the direction which she wishes them to take; and she continues these manœuvres until she has employed the whole lamina that she had separated from her body, when she takes a second, proceeding in the same manner. She gives herself no care to compress the molecules of wax which she has heaped together. She is satisfied if they adhere to each other. At length she leaves her work, and is lost in the crowd of her companions. Another succeeds and resumes the employment, then a third. All follow the same plan of placing their little masses, and if any one, by chance, gives them a contrary direction, another coming removes them to their proper place.
"The result of all these operations is a mass or little wall of wax, with uneven surfaces, five or six lines (twelfths of an inch) long, two lines high, and half a line thick, which descends perpendicularly. In this first work is no angle nor any trace of the figure of the cells. It is a simple partition in a right line without any inflection.
"The wax-makers having thus laid a foundation of a comb, are succeeded by the nurse-bees [here Huber is wrong[2]], which are alone competent to model and perfect the work. The former are the labourers, who convey the stone and mortar; the latter the masons, who work them up into the form which the intended structure requires. One of these bees now places itself horizontally on the vault (or bar-frame) of the hive, its head corresponding to the centre of the mass or wall which the wax-makers have left, and which is to form the partition of the comb into two opposite assemblages of cells; and with its mandibles (jaws), rapidly moving its head, it moulds in that side of the wall a cavity which is to form the base of one of the cells, to the diameter of which it is equal. When it has worked some minutes it departs, and another takes its place, deepening the cavity, heightening its lateral margins by heaping up the wax to right and left, by means of its teeth and fore-feet, and giving to them a more upright form. More than twenty bees successively employ themselves in this work.
[2] See remark immediately preceding the quotation.
"When arrived at a certain point, other bees begin on the yet untouched and opposite side of the mass, and commencing the bottom of two cells, are in turn relieved by others. While still engaged in this labour the wax-makers return, and add to the mass, augmenting its extent in every way, the builders again continuing their operations. After having worked the bottom of the cells of the first row into their proper forms, they polish them, and give them their finish, while others begin the outline of a new series.
"The cells themselves, or prisms, which result from the reunion and meeting of the sides, are next constructed. These are engrafted on the borders of the cavities hollowed in the mass. The bees begin them by making the contour of the bottoms, which is at first unequal, of equal height. Thus all the margins of the cells offer an uniformly level surface from their first origin, and until they have acquired their proper length. The sides are heightened in an order analogous to that which the insects follow in finishing the bottom of the cells, and the length of these tubes is so perfectly proportioned that there is no observable inequality between them."
Thus writes the great Swiss observer of bees. Without quoting at greater length from his published observations, we may give some additional particulars relating to the geometrical characters of honey-comb.
The cells of the first row laid down are pentagonal in shape. This gives them a stronger attachment to the hive than if they had had the hexagonal figure of the succeeding rows. But no form besides the six-sided prism would have answered all the conditions of the problem "how with the least expenditure of material to secure the greatest available space with the best arrangement for the purposes to be served."
Approached from the purely theoretical side, the question has been investigated by mathematicians. It requires no great acumen to determine that a hexagon of some sort is the geometrical figure which must be adopted. An equilateral triangle would make a very unsuitable abode for an insect with a nearly round body. A square cell would hardly be more convenient. A series of circles would, of course, leave interstices between them, causing a useless expenditure of space, material, time and strength. A further difficulty would arise with regard to the storage of the honey, which finds points of attachment in the angles of a hexagon, and so is less liable to run out of the cells.
The next matter then to settle is, the magnitude of the angles at which the sides of the hexagon should slope towards each other, so as to be the most advantageous. Réaumur put the problem in mathematical language before M. König, a skilful geometrician, thus:—"To determine by calculations what ought to be the angle of a hexagonal cell, with a pyramidal bottom, formed of three similar and equal rhomboid plates, so that the least matter possible might enter into its construction." The result of his investigations was that the angles of the rhombs must be 109° 26′ and 70° 34′. Cramer, professor of mathematics in the University of Geneva, also undertook the problem. His calculations, made on somewhat different principles from König's, gave for the angles 109° 28′ 16″, and 70° 31′ 44″. Maraldi, a third mathematician, assuming the equality of the angles of the trapezia forming the sides of the hexagon adjacent to the rhombs and those of the rhombs themselves, and that the solid angle at the apex of the pyramid, composed of equal obtuse angles, is precisely equal to each of the three angles at the base, also composed of three equal obtuse angles, came to the conclusion that the angles must be 109° 28′ and 70° 32′.
These three sets of results, so remarkably accordant, when we consider the minuteness of the differences between them, in figures so small as the actual honey-comb cells, show the closest correspondence to the actual measurements of the work of the bees. Maraldi found the angles of the latter to be 110° and 70°, as nearly as could be ascertained. We have dwelt at some length upon this point, because it illustrates, in a most marvellous manner, the power of that inborn faculty we call instinct, which arrives, without training, at results so precisely agreeing with those of the highest efforts of our intellectual reasonings. To the devout mind, the conclusion is inevitable that Divine Wisdom is the inspiring force which energizes the mental operations of the bees in their cell-building.
A further advantage of the actual shape of the honey-comb prisms is that, thereby, strength is combined with economy. No other form would so efficiently have carried the heavy weights constantly stored in the forms of honey, brood, and bee-bread.
The bottoms and sides of the cells are made of wax as thin as a sheet of writing-paper; but as walls of this thinness at the entrances would break down under the weight of the constantly passing insects, the margin at the opening of each cell is made four or five times thicker than the walls. Then, as the cells are lengthened, this thickness is reduced, always remaining the same, however, at the actual margins. Dr. Barclay also discovered that, though the tenuity of the divisions is so great, each, in point of fact, consists of two distinct layers agglutinated together. This gives, again, an increase of strength, as any practical builder would know who, in his "bressummers," adopts the same method of attaining lightness and power of sustaining great weights.
The actual size of the cells in a hive varies considerably, as we might expect. Without regarding those for queen-progeny, we should anticipate that those in which young drones are to be developed would be considerably larger than those prepared for workers. This is, indeed, the case. But as an abrupt change from the one kind to the other would be impossible without waste, the bees prudently graduate the difference by interposing a suitable series of intermediate sizes, whose bottoms, of course, have to depart from the normal conditions, and sometimes consist of two rhomboids and two hexagons, varying in size and form, and corresponding with four, instead of three, opposite cells. In these, stores are often found, instead of brood. If eggs are laid in them, they are generally those which will develop into males, and the space for development being smaller than usual, the drones occupying such cells are not so large as the average size.
As a rule, the hexagonal ends of twenty-seven worker cells, or nineteen drone cells, occupy a surface of one square inch. All the cells lie not quite horizontally, but sloping slightly downwards from the mouth towards the bases. This arrangement is designed to prevent the honey from easily flowing out. As the cells are filled with the liquid, the lower edge of each is first raised, and, in due time, the whole of the once open end is sealed over with a coating of wax mixed with a little propolis. This covering not only keeps the contents from running out, but prevents fermentation or candying, from contact with the air.
Each comb consists of a double layer of cells, back to back, and forming a sort of flat cake. At first this is lenticular in shape, the middle part being advanced rather more rapidly than the ends.
It is a curious fact that the bees do not, on being put into a hive, or when working in a bell-glass, begin several combs at once; but, having thoroughly laid the foundation of one, and having made some progress with this, they then start one on each side of the first, and, after a time, one on the outer side of each of the last begun. Usually, therefore, the combs hang in parallel series. If any obstruction occurs, a deviation from the normal direction takes place, but, manifest intelligence is shown in surmounting the difficulty, whatever it may be.
At first, the substance of the cells is of a dull, semi-transparent, white colour, soft, and very brittle. After a time, a yellow tinge spreads over the comb, and, with age, this hue deepens to brown, and if some years old, becomes almost black. The colour, therefore, furnishes a tolerably safe guide as to the age of comb. The darkening seems due, partly to a chemical change from contact with the air, but still more to the constant traffic of the bees over it, and its getting smeared with dirt and propolis.
It occasionally happens that, owing to a great in-flow of honey, the weight of the combs endangers their security, and the bees, seeing the danger of their breaking down, resort to a most clever method of rendering their treasures safe. Gnawing away a small part of the topmost row of the combs on one side, they lay a broader foundation, and then, with a strongly glutinous mixture of wax and propolis, they fasten afresh the upper cells to their points of attachment. Having completed one side, they then proceed in the same way with the other, till they are satisfied of the firmness of the whole structure.
Again, if the supply of food outruns the capacity of their store-houses as first made, they will often lengthen the cells, till, especially in the case of supers, they reach the length of even two inches—more than twice the normal size.
The queen-cells are remarkably distinguished from those for workers or drones, in respect to size, direction, shape, and amount of material. They occupy at least as much space as half a dozen ordinary cells. They are directed downwards, instead of lying horizontally. They are irregularly oval or pyriform in shape, and are made up of a sort of mosaic of wax, which material, so sparingly used elsewhere, seems lavished on the royal nurseries. The reasons for this are, probably, to secure the young queens from danger while passing through the larval and pupal conditions, and to keep up the warmth necessary for their more rapid development. Wax being a very bad conductor of heat, the thick walls prevent the chilling of the brood, and, at the same time, allow of considerable clustering of nurse-bees, and consequent generation of warmth, without the danger of the cells being broken down by the pressure.
Bees-wax forms a not unimportant article of commerce. From Germany, Greece, Cyprus, and still more largely from North America, we derive what is needed to make up the deficiency in our home production of it. Its uses are numerous. For household purposes, especially for polishing furniture, for some varnishes and unguents, for candles and matches, for modelling, particularly in dentistry, it is consumed in great quantities. Since the introduction of paraffin and similar substances for lighting purposes, the amount used for candles has diminished, though the demand for it in other directions does not appear to have fallen off. Bee-keepers now use it greatly for "foundation-comb."