Origin of Species by Charles Darwin
Chapter VIII. Instinct
Cell-making instinct of the hive-bee
I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wax in their construction. It has been remarked that a skilful workman, with fitting tools and measures, would find it very difficult to make cells of wax of the true form, though this is effected by a crowd of bees working in a dark hive. Granting whatever instincts you please, it seems at first quite inconceivable how they can make all the necessary angles and planes, or even perceive when they are correctly made. But the difficulty is not nearly so great as at first appears: all this beautiful work can be shown, I think, to follow from a few simple instincts.
I was led to investigate this subject by Mr. Waterhouse, who has shown that the form of the cell stands in close relation to the presence of adjoining cells; and the following view may, perhaps, be considered only as a modification of his theory. Let us look to the great principle of gradation, and see whether Nature does not reveal to us her method of work. At one end of a short series we have humble-bees, which use their old cocoons to hold honey, sometimes adding to them short tubes of wax, and likewise making separate and very irregular rounded cells of wax. At the other end of the series we have the cells of the hive-bee, placed in a double layer: each cell, as is well known, is an hexagonal prism, with the basal edges of its six sides bevelled so as to join an inverted pyramid, of three rhombs. These rhombs have certain angles, and the three which form the pyramidal base of a single cell on one side of the comb, enter into the composition of the bases of three adjoining cells on the opposite side. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humble-bee, we have the cells of the Mexican Melipona domestica, carefully described and figured by Pierre Huber. The Melipona itself is intermediate in structure between the hive and humble bee, but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells, in which the young are hatched, and, in addition, some large cells of wax for holding honey. These latter cells are nearly spherical and of nearly equal sizes, and are aggregated into an irregular mass. But the important point to notice is, that these cells are always made at that degree of nearness to each other that they would have intersected or broken into each other if the spheres had been completed; but this is never permitted, the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Hence, each cell consists of an outer spherical portion, and of two, three, or more flat surfaces, according as the cell adjoins two, three or more other cells. When one cell rests on three other cells, which, from the spheres being nearly of the same size, is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. As in the cells of the hive-bee, so here, the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. It is obvious that the Melipona saves wax, and what is more important, labour, by this manner of building; for the flat walls between the adjoining cells are not double, but are of the same thickness as the outer spherical portions, and yet each flat portion forms a part of two cells.
Reflecting on this case, it occurred to me that if the Melipona had made its spheres at some given distance from each other, and had made them of equal sizes and had arranged them symmetrically in a double layer, the resulting structure would have been as perfect as the comb of the hive-bee. Accordingly I wrote to Professor Miller, of Cambridge, and this geometer has kindly read over the following statement, drawn up from his information, and tells me that it is strictly correct:-
If a number of equal spheres be described with their centres placed in two parallel layers; with the centre of each sphere at the distance of radius x sqrt(2) or radius x 1.41421 (or at some lesser distance), from the centres of the six surrounding spheres in the same layer; and at the same distance from the centres of the adjoining spheres in the other and parallel layer; then, if planes of intersection between the several spheres in both layers be formed, there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs; and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee. But I hear from Professor Wyman, who has made numerous careful measurements, that the accuracy of the workmanship of the bee has been greatly exaggerated; so much so, that whatever the typical form of the cell may be, it is rarely, if ever, realised.
Hence we may safely conclude that, if we could slightly modify the instincts already possessed by the Melipona, and in themselves not very wonderful, this bee would make a structure as wonderfully perfect as that of the hive-bee. We must suppose the Melipona to have the power of forming her cells truly spherical, and of equal sizes; and this would not be very surprising, seeing that she already does so to a certain extent, and seeing what perfectly cylindrical burrows many insects make in wood, apparently by turning round on a fixed point. We must suppose the Melipona to arrange her cells in level layers, as she already does her cylindrical cells; and we must further suppose, and this is the greatest difficulty, that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres; but she is already so far enabled to judge of distance, that she always describes her spheres so as to intersect to a certain extent; and then she unites the points of intersection by perfectly flat surfaces. By such modifications of instincts which in themselves are not very wonderful--hardly more wonderful than those which guide a bird to make its nest--I believe that the hive-bee has acquired, through natural selection, her inimitable architectural powers.
But this theory can be tested by experiment. Following the example of Mr. Tegetmeier, I separated two combs, and put between them a long, thick, rectangular strip of wax: the bees instantly began to excavate minute circular pits in it; and as they deepened these little pits, they made them wider and wider until they were converted into shallow basins, appearing to the eye perfectly true or parts of a sphere, and of about the diameter of a cell. It was most interesting to observe that, wherever several bees had begun to excavate these basins near together, they had begun their work at such a distance from each other that by the time the basins had acquired the above stated width (i.e. about the width of an ordinary cell), and were in depth about one sixth of the diameter of the sphere of which they formed a part, the rims of the basins intersected or broke into each other. As soon as this occurred, the bees ceased to excavate, and began to build up flat walls of wax on the lines of intersection between the basins, so that each hexagonal prism was built upon the scalloped edge of a smooth basin, instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells.
I then put into the hive, instead of a thick, rectangular piece of wax, a thin and narrow, knife-edged ridge, coloured with vermilion. The bees instantly began on both sides to excavate little basins near to each other, in the same way as before; but the ridge of wax was so thin, that the bottoms of the basins, if they had been excavated to the same depth as in the former experiment, would have broken into each other from the opposite sides. The bees, however, did not suffer this to happen, and they stopped their excavations in due time; so that the basins, as soon as they had been a little deepened, came to have flat bases; and these flat bases, formed by thin little plates of the vermilion wax left ungnawed, were situated, as far as the eye could judge, exactly along the planes of imaginary intersection between the basins on the opposite side of the ridge of wax. In some parts, only small portions, in other parts, large portions of a rhombic plate were thus left between the opposed basins, but the work, from the unnatural state of things, had not been neatly performed. The bees must have worked at very nearly the same rate in circularly gnawing away and deepening the basins on both sides of the ridge of vermilion wax, in order to have thus succeeded in leaving flat plates between the basins, by stopping work at the planes of intersection.
Considering how flexible thin wax is, I do not see that there is any difficulty in the bees, whilst at work on the two sides of a strip of wax, perceiving when they have gnawed the wax away to the proper thinness, and then stopping their work. In ordinary combs it has appeared to me that the bees do not always succeed in working at exactly the same rate from the opposite sides; for I have noticed half-completed rhombs at the base of a just-commenced cell, which were slightly concave on one side, where I suppose that the bees had excavated too quickly, and convex on the opposed side where the bees had worked less quickly. In one well-marked instance, I put the comb back into the hive, and allowed the bees to go on working for a short time, and again examined the cell, and I found that the rhombic plate had been completed, and had become perfectly flat: it was absolutely impossible, from the extreme thinness of the little plate, that they could have effected this by gnawing away the convex side; and I suspect that the bees in such cases stand in the opposed cells and push and bend the ductile and warm wax (which as I have tried is easily done) into its proper intermediate plane, and thus flatten it.
>From the experiment of the ridge of vermilion wax we can see that, if the bees were to build for themselves a thin wall of wax, they could make their cells of the proper shape, by standing at the proper distance from each other, by excavating at the same rate, and by endeavouring to make equal spherical hollows, but never allowing the spheres to break into each other. Now bees, as may be clearly seen by examining the edge of a growing comb, do make a rough, circumferential wall or rim all round the comb; and they gnaw this away from the opposite sides, always working circularly as they deepen each cell. They do not make the whole three-sided pyramidal base of any one cell at the same time, but only that one rhombic plate which stands on the extreme growing margin, or the two plates, as the case may be; and they never complete the upper edges of the rhombic plates, until the hexagonal walls are commenced. Some of these statements differ from those made by the justly celebrated elder Huber, but I am convinced of their accuracy; and if I had space, I could show that they are conformable with my theory.
Huber's statement, that the very first cell is excavated out of a little parallel-sided wall of wax, is not, as far as I have seen, strictly correct; the first commencement having always been a little hood of wax; but I will not here enter on details. We see how important a part excavation plays in the construction of the cells; but it would be a great error to suppose that the bees cannot build up a rough wall of wax in the proper position--that is, along the plane of intersection between two adjoining spheres. I have several specimens showing clearly that they can do this. Even in the rude circumferential rim or wall of wax round a growing comb, flexures may sometimes be observed, corresponding in position to the planes of the rhombic basal plates of future cells. But the rough wall of wax has in every case to be finished off, by being largely gnawed away on both sides. The manner in which the bees build is curious; they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell, which will ultimately be left. We shall understand how they work, by supposing masons first to pile up a broad ridge of cement, and then to begin cutting it away equally on both sides near the ground, till a smooth, very thin wall is left in the middle; the masons always piling up the cut-away cement, and adding fresh cement on the summit of the ridge. We shall thus have a thin wall steadily growing upward but always crowned by a gigantic coping. From all the cells, both those just commenced and those completed, being thus crowned by a strong coping of wax, the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls. These walls, as Professor Miller has kindly ascertained for me, vary greatly in thickness; being, on an average of twelve measurements made near the border of the comb, 1/352 of an inch in thickness; whereas the basal rhomboidal plates are thicker, nearly in the proportion of three to two, having a mean thickness, from twenty-one measurements, of 1/229 of an inch. By the above singular manner of building, strength is continually given to the comb, with the utmost ultimate economy of wax.
It seems at first to add to the difficulty of understanding how the cells are made, that a multitude of bees all work together; one bee after working a short time at one cell going to another, so that, as Huber has stated, a score of individuals work even at the commencement of the first cell. I was able practically to show this fact, by covering the edges of the hexagonal walls of a single cell, or the extreme margin of the circumferential rim of a growing comb, with an extremely thin layer of melted vermilion wax; and I invariably found that the colour was most delicately diffused by the bees--as delicately as a painter could have done it with his brush--by atoms of the coloured wax having been taken from the spot on which it had been placed, and worked into the growing edges of the cells all round. The work of construction seems to be a sort of balance struck between many bees, all instinctively standing at the same relative distance from each other, all trying to sweep equal spheres, and then building up, or leaving ungnawed, the planes of intersection between these spheres. It was really curious to note in cases of difficulty, as when two pieces of comb met at an angle, how often the bees would pull down and rebuild in different ways the same cell, sometimes recurring to a shape which they had at first rejected.
When bees have a place on which they can stand in their proper positions for working--for instance, on a slip of wood, placed directly under the middle of a comb growing downwards, so that the comb has to be built over one face of the slip--in this case the bees can lay the foundations of one wall of a new hexagon, in its strictly proper place, projecting beyond the other completed cells. It suffices that the bees should be enabled to stand at their proper relative distances from each other and from the walls of the last completed cells, and then, by striking imaginary spheres, they can build up a wall intermediate between two adjoining spheres; but, as far as I have seen, they never gnaw away and finish off the angles of a cell till a large part both of that cell and of the adjoining cells has been built. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two just-commenced cells, is important, as it bears on a fact, which seems at first subversive of the foregoing theory; namely, that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal; but I have not space here to enter on this subject. Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells, if she were to work alternately on the inside and outside of two or three cells commenced at the same time, always standing at the proper relative distance from the parts of the cells just begun, sweeping spheres or cylinders, and building up intermediate planes.
As natural selection acts only by the accumulation of slight modifications of structure or instinct, each profitable to the individual under its conditions of life, it may reasonably be asked, how a long and graduated succession of modified architectural instincts, all tending towards the present perfect plan of construction, could have profited the progenitors of the hive-bee? I think the answer is not difficult: cells constructed like those of the bee or the wasp gain in strength, and save much in labour and space, and in the materials of which they are constructed. With respect to the formation of wax, it is known that bees are often hard pressed to get sufficient nectar; and I am informed by Mr. Tegetmeier that it has been experimentally proved that from twelve to fifteen pounds of dry sugar are consumed by a hive of bees for the secretion of a pound of wax; so that a prodigious quantity of fluid nectar must be collected and consumed by the bees in a hive for the secretion of the wax necessary for the construction of their combs. Moreover, many bees have to remain idle for many days during the process of secretion. A large store of honey is indispensable to support a large stock of bees during the winter; and the security of the hive is known mainly to depend on a large number of bees being supported. Hence the saving of wax by largely saving honey, and the time consumed in collecting the honey, must be an important element of success any family of bees. Of course the success of the species may be dependent on the number of its enemies, or parasites, or on quite distinct causes, and so be altogether independent of the quantity of honey which the bees can collect. But let us suppose that this latter circumstance determined, as it probably often has determined, whether a bee allied to our humble-bees could exist in large numbers in any country; and let us further suppose that the community lived through the winter, and consequently required a store of honey: there can in this case be no doubt that it would be an advantage to our imaginary humble-bee if a slight modification of her instincts led her to make her waxen cells near together, so as to intersect a little; for a wall in common even to two adjoining cells would save some little labour and wax. Hence, it would continually be more and more advantageous to our humble-bees, if they were to make their cells more and more regular, nearer together, and aggregated into a mass, like the cells of the Melipona; for in this case a large part of the bounding surface of each cell would serve to bound the adjoining cells, and much labour and wax would be saved. Again, from the same cause, it would be advantageous to the Melipona, if she were to make her cells closer together, and more regular in every way than at present; for then, as we have seen, the spherical surfaces would wholly disappear and be replaced by plane surfaces; and the Melipona would make a comb as perfect as that of the hive-bee. Beyond this stage of perfection in architecture, natural selection could not lead; for the comb of the hive-bee, as far as we can see, is absolutely perfect in economising labour and wax.
Thus, as I believe, the most wonderful of all known instincts, that of the hive-bee, can be explained by natural selection having taken advantage of numerous, successive, slight modifications of simpler instincts; natural selection having, by slow degrees, more and more perfectly led the bees to sweep equal spheres at a given distance from each other in a double layer, and to build up and excavate the wax along the planes of intersection. The bees, of course, no more knowing that they swept their spheres at one particular distance from each other, than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates; the motive power of the process of natural selection having been the construction of cells of due strength and of the proper size and shape for the larvae, this being effected with the greatest possible economy of labour and wax; that individual swarm which thus made the best cells with least labour, and least waste of honey in the secretion of wax, having succeeded best, and having transmitted their newly-acquired economical instincts to new swarms, which in their turn will have had the best chance of succeeding in the struggle for existence.