Art. XLV.—On the Construction of the Comb of the Hive-bee.
[Read before the Wellington Philosophical Society, 4th September, 1895.]
During the past autumn, whilst shifting my bee-boxes, I disturbed a couple of hives to such an extent that the bees deserted them. These boxes were really double hives (two boxes to each hive), with a hole, 3 ¼in. square, leading from the lower to the upper box. I had consequently a good opportunity of carefully examining the deserted comb in the lower boxes, which I will name A and B. To my surprise I noticed that the square hole in the top of box A had been bridged across by two walls of comb, whilst in box B a combbridge leading straight across the hole had only just been begun. This was all the comb in box B.
Box A, it will be noticed, is almost full of comb, lines of it stretching from side to side, and two of these across the square hole, proving that, in both instances, the bees, having apparently satisfied themselves of the dimensions of the holes, ignored them completely, and relied upon the bridging-powers of their comb-construction to span the distances. The fact that in box B the new line of comb (which, when complete, would stretch from side to side of the box) had been begun directly at the aperture, and the bridge of comb at once thrown straight across, almost reaching to the other side, shows how thoroughly the bees recognised the obstacle, seeing that they had the whole of the box at their disposal in which to start operations.
Darwin makes no reference to these honeycomb bridges in his “Origin of Species.” All that great writer does is to strive his utmost to show that the comb of the hive-bee arose in accordance with his great theory of natural selection. He enters very fully into the cell-making instinct of the hive-bee. He cites other authorities to support his theory, which, if wrong in this instance, is certainly wrong throughout his
whole work. So completely do I disagree with the theory as explaining the origin of species, so certain am I that there is another force, energy, or intelligence in nature far superior to it (which I have named “a common vital force”)—an intelligence that acts equally upon all living things throughout the whole universe we see around us—that I have determined to pin Darwin down to his own words in regard to this one matter of the cell-making instinct of bees, and to show, to the best of my ability, how the theory of natural selection fails to explain anything at all regarding bee-life—why it should, in some respects, resemble the busy life of a human community, or this wonderful specimen of what is called “instinct” in the construction of honeycomb. Oftentimes have my friends said to me, “Look how well Darwin explains his principle of evolution by natural selection in the superiority of the hive-bee cell to that of the humble-bee!” I propose now to ask members whether he tells us anything at all in this one special matter.
The following is Darwin's conclusion to his section (cap. viii., p. 227, “Origin of Species”) : “Thus, 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 larvæ, 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.”
Nothing can be plainer or more emphatic than these words as explaining the theory of natural selection. I have italicised certain words in order not only to show how mistaken this conclusion was, but also to emphasize the existence of this constant guiding vital intelligence, of which Darwin took no account. There is no reference to the words, “vital intelligence or force” in Darwin's index. It is of no moment whether he himself compiled the index or the index was compiled for him :
I should certainly think he supervised it. But it is quite plain that so little was thought of these two words that they were omitted from the index. In his conclusion Darwin certainly continues with the words, “the motive-power of the process of natural selection,” as being sufficient to account for the cell-construction of the hive-bee; but no person has yet granted natural selection to be a motive-power. Every one admits that vital intelligence is a motive-power, because we all experience it, although we do not know what it really is; and we can all further admit that natural selection is one method by which that intelligence acts, subject, however, to the higher law of progressive adaptation of species. That is to say, that each species, like a magic puzzle, has in itself the power to change, to adapt itself, to build cell on cell, by a thousand thousand different modifications, so as to enable it to suit itself to new environments. I pointed this out in section iii. of my paper—“Potentiality of Divergence.”*
The bridging principle of construction in the comb of the hive-bee can be seen at a glance in both boxes on the table. There is no simple cell-construction in the bold manner in which the bees throw their comb straight across the two holes —no simple “sweeping of equal spheres from respective distances.” Again, it will be observed that the walls of comb stand at certain distances from each other—never less than three lines, but usually six lines—that being, I suppose, as far as the bees can reach between wall and wall of comb. Surely the bees know what they are doing when rigidly keeping the different lines of comb at these stated distances. For it will be observed that, no matter how the bases of the hanging walls of comb start, whether in straight or diagonal lines, each single wall rarely approaches a neighbouring wall within three lines. These bases, indeed, can start anywhere in the box, because the bees evidently possess two separate pieces of knowledge amongst many others: (1) The average length of the rhombus or cell to accommodate the larvæ; (2) the distance to be observed between the walls of growing comb. Possessing these two pieces of knowledge, which Darwin does not credit the bees with, as well as the “instinct of the sweeping of spheres” to form the cells, which he does credit them with, the bees can fill in a space anywhere in the box. The result always is that a box becomes filled with walls of live comb at stated distances apart, and the two layers of cells of fairly equal lengths. But should a growing wall of comb come in contact with a neighbouring wall, as sometimes happens, that contact is immediately stopped, and the point of contact forms a stay. It will be observed that these stays
[Footnote] * Trans. N.Z. Inst., 1893, p. 611.
are numerous round the sides of the box. This is evidently to strengthen the walls of comb and keep them firm, the stays rarely impeding the free passing of the bees round the hive.
In Article lxxi. of our Transactions of 1893, “Spiders as Engineers,” I pointed out how beautifully spiders stayed their webs, and showed, moreover, that some of our own suspension-bridges (notably the one in Hobson Street, Wellington) are stayed exactly in a similar manner. Now, if any person closely examines box A he will observe somewhat similar stays all round the box, but rarely between the walls of comb. What, then, is the intelligence that guides insects and animals to stay their constructions in this way? Wherein does the theory of natural selection account for it? Does nature, under that theory, thin out all the variations of the different species until only those survive which know how to adopt this principle of staying? For that is the constant argument: “Only those survive that have been naturally selected to survive.” Now, granting this argument, under the higher law of progressive adaptation, how does the theory account for such widely-different species as man, spiders, and bees using almost the same principle in staying their widely-different structures? For, whether the stay is made of wax, web, or iron, there the principle is all the same. To say that “Similarity of object leads to similarity of means” implies that there is in nature a previously-existing method or means to a particular end for all species.
Furthermore, it will be observed that the walls of comb run fairly straight: so that bees know how to make fairly straight lines, as well as how to make bridges. I do not say that every wall is absolutely straight—there are curves in some—but the tendency is to run straight lines. Of the three centre lines of comb in box A, the bees, it will be observed, had a guide for two of the walls in the little gap where the two boards forming the top of the box should have closely met. The third wall, of course, followed the centre wall near the gap after it had been constructed.
The reason for the diagonal walls is not clear. I have seen boxes of comb with all the walls fairly parallel and straight excepting in one corner, where the beautiful white comb forming the queen's home is built. This comb is usually built diagonally, perhaps for purposes of easier defence should an enemy invade the hive. The fact of aiming at straight lines at all, even in the diagonal walls, shows a bee's further knowledge, which Darwin ignored. I do not for one moment mean to say that the principle of natural or artificial selection is absolutely non-existent in nature. What I mean is that in the higher law of progressive adaptation of species, natural selection plays, as I have said, only a minor part.
I may be allowed here to digress for one moment to give an example of progressive development which forms one phase of progressive adaptation as distinct from the theory of natural selection. It was formerly supposed that a very minute crustaceous animal inhabiting the open sea, named Zoea, was totally distinct from the genus Megalopa, which again was supposed to be totally distinct from every known genus of Crustacea. Gosse very clearly points out that “These conclusions were set aside by the brilliant discovery of Dr. Vaughan Thompson that Zoea and Megalopa were the same animal in different stages of existence; and that, moreover, both were but the early states of well-known and familiar forms of larger Crustacea, which therefore undergo a metamorphosis as complete as that by which the caterpillar changes to a chrysalis and the chrysalis to a butterfly, and in every essential point parallel to it. In the Cove of Cork Dr. Thompson met with a considerable number of Zoeas, which he kept in captivity. Some of these changed into the Megalopa form, which in turn changed to the most abundant of all our larger Crustacea, the common shore-crab (Carcinus mœnas). Thus, in its progress from the egg to its final development, the crab was forced to pass through two temporary conditions, which had previously been regarded as types not of genera only, but of different families, and both strikingly dissimilar from the group to which, in its perfect state, it belongs.”
Here we have an example of progressive development which quite puzzled Darwin himself under his own theory. But we know of many other instances of peculiar stages of development in nature to which the theory of natural selection does not at all apply, and I propose to refer to them later on.
But let us proceed. It will be observed that the sides of the comb-walls—the ends of the layers—are finished off almost in perfect planes. Pass the fingers carefully over them, and, no matter whether the distances between the walls be three, five, seven, or nine lines (for my non-scientific hearers I may explain that twelve lines go to the inch), it will be seen how beautifully the bees know when to cease the length of the cell or rhomb. (In geometry a rhomb or rhombus is an oblique-angled equilateral parallelogram.) Here is a further piece of knowledge on the bees' part of which Darwin took no account. I may explain that it is by the dexterous use of their stings that the bees finish off and cap their cells, injecting a minute portion of formic acid into the honey as the cell is filled and closed. This acid is really the poison of their stings, and it imparts to the honey its peculiar flavour and keeping-qualities. The sting is an exquisitely-contrived little trowel, and it greatly helps in giving the plane surface to the sides of the layers. I may further be allowed to say that to describe the cylindrical
tongue of the bee, which laps up the honey, almost exceeds, according to Suammerdam and Gosse, the utmost efforts of human knowledge.
Now, hold a piece of comb up to the light, or break off a piece. It will be observed that the cells forming each side of the comb-wall—the two layers—do not start from the same base. The hexagon is not continued straight through the wall, but is broken in the centre, forming the basal rhombic plate. This I take to be the most marvellous work of construction in the comb; for here the bees know exactly how to break the joint, for the special purpose, I suppose, of giving strength to the comb-wall and to the two opposite cells.
There is no “blind sweeping of equal spheres at stated distances” in this breaking of the joint, for the bees know exactly how to place and plane the basal plates and angles out of the wax so as to perform this most delicate and wonderful principle of construction. Therefore, whatever the guiding principle of construction in nature may be which controls such a work, the principle of natural selection is not within a thousand miles of it. We can admit, for the sake of argument, that there may be a principle of natural selection—I do not for one moment say there is—between the simple cocoons of the humble-bee and the cells of the hive-bee, of which the cell of the Mexican Melipona domestica is the intermediate stage. This is what we are asked to admit. Pierre Huber, however, who has carefully described and figured the cell of Melipona domestica, calls it a “gross imitation” of the three-sided pyramidal base of the cell of the hive-bee. Darwin ignored these words completely, and made use of Huber's name as a support for his theory. But what has this very short series of natural-selection stages—(so very short a one, with so few examples, and these so very uncertain, that I am completely surprised Darwin did not himself candidly admit his want of proof, in place of taking it for granted that we should accept his theory as a matter of course : he, moreover, drawing in the name of Professor Nyman to support his assumption by the statement that “the accuracy of the workmanship of the bee has been greatly exaggerated”)—what even has this short series to do with breaking the joint at the base of each cell? Surely the bases might have been equal in the layers for the starting of the sides of each rhombus! But they are not so, and the bees know that they are not to be so, just as surely as the horse-bot knows that the safest place to deposit its eggs is just beneath the horse's chin. Evolutionists will, of course, say that the horse-bot has been naturally selected to do this. I propose to expose the fallacy of such an argument later on.
It is not necessary for me to go into the actual details of the cell-making, the gathering of the honey and the secretion
of the wax, its deposit, working and planing off into hexagonal or basal plates. An observer must watch the bees at work in a glass hive, or read any bee-book upon the subject. The sets of combs on the bees' hind feet for scraping up the pollen, and the little baskets or paniers on the tibia joints immediately above these combs for carrying it to the hive, are so wonderful in their construction that I can only marvel at this one display of Divine intelligence. My mind positively recoils from ascribing it to any blind principle of natural selection. I will admit a slight “sweeping of equal spheres at given distances,” because the bees have to work in the dark by feel and sound, and to economize space. There are, I think, one or two other senses than ours amongst bees, of which we at present know little or nothing. In referring to this point Gosse says, “The comparative moisture or dryness of the atmosphere, delicate changes in its temperature, in its density; the presence of gaseous exhalations; the proximity of solid bodies indicated by subtle vibrations of the air; the height above the earth at which flight is performed, measured basometrically; the various electrical conditions of the atmosphere; and perhaps many other physical diversities which cannot be classed under sight, sound, smell, taste, or touch, and which may be altogether unappreciable, and therefore altogether inconceivable, by us.” To which I may add now the “sembling” of insects, more especially that of the oak egger-moth (Lasiocampa quercus). But there are a hundred marvels of vital action, energy, or intelligence connected with a bee-hive—such as the bees always turning their faces to the queen (just as man does to his queen) as she moves through the hive; their choice of a queen, and all the struggles consequent thereupon; their building a palace of beautiful comb for the queen, and its strategical defence; their killing off the drones or superfluous bees before the winter sets in; their crooning the night before swarming, just as men and women croon and cry when they have to leave their parent homes; their swarming, and their hanging about in the vicinity of the parent hive for two or three days in order to see whether man will put them in a box and place them near their birthplace; their harmlessness whilst waiting about, and the ease by which an experienced bee-taker can sweep them with his bare hand into their new hive, &c. All these things rest upon the immutable principles of a guiding vital intelligence, from which natural selection is as far removed as the sun from the earth. All this is done by what we have perhaps mistakenly named “instinct.”
But closely examine the new swarm when clustering upon a bough—there is little danger, as bees rarely sting during swarming time—it will be observed that the individual bees are constantly on the move; those on the inside coming to
the outside of the cluster, and those upon the outside going inwards to relieve those holding on to the bough, and to give them a spell from bearing the heavy weight of the cluster. The young bees, new to life, know that it is their bounden duty to do this, and not to go flying about seeking honey. Will any person for one moment say that the bees have been taught to do this sensible work by the principle of natural selection ?—that only those bees have survived which have known how to swarm properly? Or will they not admit with me that there is some guiding energy or intelligence which tells the young bees how to swarm now, as it has told them for a million years, and just as it tells them how to plane and shape the basal rhombic plates of their cells?
It will be noticed in box A that the hanging wall of comb grows downwards until it reaches the bench upon which the box rests within a distance of about ½in. Pass a straightedge over the box and this space will at once be seen. This space is left for ventilation and room in the hive. The bees have to think of proper ventilation just as we have. On very warm, calm days a certain number stand at the entrance to their hives and convert themselves into miniature air-fans by rapid vibration of their wings. No doubt the comb-walls are constructed to afford a free ventilating-space. The cell therefore lengthens outwards as the comb-wall descends, the wall itself usually having a sharp edge, which is the first forming of the basal rhombic plates. But, no matter how these wonderful basal plates start, the rhomb or cell very slightly curves outwards from them, and lengthens to its proper position in the layer. There appears to be only one set of basal plates in the comb-wall, even when one side of the wall is lengthened to three times the ordinary length of the cell so as to form a stay. Of course, there cannot well be more, otherwise there would be a closed and vacant cell in the centre of the wall. The bees know quite well how and when to make their cells of different lengths as circumstances require. In this a most marvellous intelligence is displayed, the cells having to vary by gradation from the ordinary to three times their ordinary length. The mere fact of lengthening the cell into different lengths as required shows that there is a guiding principle at work far removed from the mere “blind sweeping of equal spheres at stated distances.” The bees work from cell to cell, from base and angle, with mathematical precision, leaving the whole wall a perfectly-finished work. The marvel lies in this continuity of simplicity in the angular construction, proving undoubtedly to any unprejudiced mind that there is no blindness whatever in the bees' mode of working.
But can it be said that the hexagonal cell is the result
of a sweeping of an equal sphere at all? Examine the cell. It is six-sided, almost mathematically correct. The basal plates of the rhomb do not form a plane, but contain three faces or oblique angles too, so that I cannot see where the “sweeping of equal spheres” finds place in this construction. They may start the cell by sweeping a sphere, but there is no sphere left by the time the cell is finished. The bees measure the cell-distances in the layer by the size of their own bodies; and then the principle of construction is hexagonal and not circular. The cell of the humble-bee is circular; that of Melipona domestica circular, and oftentimes a “gross imitation of the cell of the hive-bee.” But can any person say that the cell of the hive-bee is not the proper cell, devised and perfected by nature, and that the humble-bee cell and that of Melipona are only variations? As a rule, it is the humble-bee that drives the hive bee away. What ground, then, has Darwin for applying his theory to this construction at all? Do his instances prove his argument? Can they not be read entirely the other way—namely, that they are but degraded variations of the proper cell? And if we find that Darwin has been absolutely mistaken in applying his theory to this one insect, what value are we to attach to his other instances of proof? The cell of the humble-bee ought to be far and away superior to the cell of the hive-bee, but it is not so.
Darwin, moreover, names it as “the most wonderful of all known instincts, that of the hive-bee.” In my opinion he is wrong. The ant shows a far more wonderful display of what is called “instinct.” I shall refer later on to the ant. The bee, ant, spider, and man show similar knowledge of exactly similar subjects. To say that the bee works without this knowledge, but simply owing to “blind evolution by natural selection, having taken advantage of numerous successive slight modifications of simpler instincts” (which modifications and instances are not given), is to me a proposition quite untenable.
A hive swarms. A certain number of the older bees may, or may not, accompany the queen and the young bees. These may or may not teach the new swarm how to begin their labours and use their little planes—like our carpenter's planes —to level off and reduce to a uniform thickness the walls of their cells. And let it be noted how beautiful and perfect the wall of the hexagon is—never breaking into an adjoining cell, but a perfectly watertight compartment for holding its store of honey, food, or young larvæ. In my opinion there is no necessity for the young brood to be taught how to go to work any more than it is taught how to swarm. In the realm of nature throughout the whole universe we see around us, certain common vital laws rule. A young hive-bee goes to work with its
little tools and builds its cell in the most cheery confidence, because it knows that food in flowers has been provided for it, and because it has been designed to build the cell exactly in this special manner to store its food. It has been as much designed to do this as it has been designed to fertilise plants. Will any person who objects to the word “designed” kindly explain how it is that numerous species of plants depend for existence and propagation almost entirely upon the visits of bees? We know of no other principle of construction that will hold so much liquid so well and in so small a space as that of the cell of the hive-bee. There is the cell of the mason-wasp, which is a very wonderful structure too, although oftentimes a great nuisance to us in New Zealand. But this cell is round or oblong, with no oblique angles—a much more simple construction than that of the hive-bee. Moreover, it is built of clay, not of wax. How is the doctrine of natural selection to explain this one vast difference of material in exactly a similar operation ? True, its whole purpose is different, but nevertheless a cell is built.*.
But after the swarm has been taken, away the young bee flies. It may have to fly a mile or more; but it brings back its nectar, unerringly selects its new home out of a row of perhaps twenty boxes, and sets to work just as I suppose its ancestors have worked throughout all time.
The dividing wall of the two layers of cells are named “basal rhombic plates.” I cannot define more than three plates with clearly-formed and beautiful oblique angles. Yet in his conclusion Darwin says, “the bees, of course, no more knowing that they swept their spheres at one particular distance from each other than they knew what are the several angles of the hexagonal prisms and of the basal rhombic plates.” Now, I ask any one really desirous of testing these statements—(and, not-withstanding the weight of authority Darwin makes a point of always bringing to support his arguments, I hold that every one of his statements requires the most careful testing and verifying)—to look at these basal plates of the cell and ask himself whether the bee knew what it was doing or whether it did not? If not, if it only acts from the blind principle of evolution and natural selection that is immediately afterwards referred to in the conclusion, why is it that each basal plate is
[Footnote] * The mason-fly, by some method I cannot explain, stupefies, either by injecting a fluid or paralysing certain nerves, the spider which it has entombed in the cell with its egg. This stupefaction lasts a period of two to four months. The larvæ of the fly hatch out and feed upon the beautifully-preserved body of the spider. I have often thought that this process of stupefaction should be carefully investigated, in order to see whether nature has not in this matter shown us an example whereby we might preserve our meat.
exactly alike, and as mathematically correct as a spider's web? What is it that keeps these angles uniform right through the comb-wall ?—for a well-constructed oblique angle is not a mere blind sweeping of a sphere.
To give Darwin every credit, which I naturally wish to give to so great a writer, I will say this: that, if this instance of natural selection in regard to the hive-bee cell he has given us offers any proof of his theory to scientific minds (I regret it does not to mine), then at best it is but an exemplification of the law of progressive adaptation of species. The hive-bee, the humble-bee, and Melipona domestica are each useful for its particular work. Red-clover, for instance, in New Zealand could not be fertilised until we introduced the humble-bee. It is also said in Canterbury that the humble-bee in some places takes all the honey from the flowers, leaving little or none for the hive-bee. The cell of the humble-bee should there-fore be far superior to the cell of the hive-bee. But Darwin places the cell of the humble-bee at the bottom of the scale, and most unmistakably says that Melipona domestica and the hive-bee cells have been naturally selected from it. To say that the humble-bee is evolved by natural selection from the hive-bee, or vice versa, or that the hive-bee cell is naturally selected from the Mexican-bee cell, looks to me quite absurd, even from Darwin's own proof. As Pierre Huber distinctly says, “The Mexican-bee cell looks like a gross imitation of a portion of the hive-bee cell.” Moreover, we have in New Zealand many native bees which build simple single cells in the ground for storing their food. The humble-bee cell is almost a clay cell. Wasps in Europe build their nests of clay. But all these are quite different structures to the finished cell of wax of the hive-bee. Again, even in the hive-bee cells there is no blind sweeping of equal spheres, seeing that the cells for the queen-bees are considerably larger than those for the common bees of the hive, and are also differently constructed.*
[Footnote] * I attach the following description of the cell-formation and work of the queen-bee as bearing upon the question. I regret my inability to give the author's name: “The province and occupation of the queen-bee consist in laying the eggs from which originate the prodigious multitudes that people a hive. Every bee in the community is apparently aware of this fact, and consequently treats her with due respect, even to the extent of never turning its back upon her until, the hive being overcrowded and a new queen having been made, a swarm is thought necessary, when all respect disappears, and, should she show the least reluctance, she is forced out to seek new quarters with other emigrants. The creation of a queen is one of the greatest wonders of that most wonderful of insect communities—a hive of bees; for no sooner does the old queen die, or the members of the community become convinced that they are overcrowded, and that a swarm is necessary, than they begin to build one or more queen-cells, which are utterly unlike the well-known hexagonal cells in which honey is stored or the brood of either workers or drones is reared, being in shape and size not unlike an acorn. In each of these, if more than one, either a worker-egg—worker- and drone-eggs being dissimilar, and laid in different comb—or a worker and worker-larva not more than three days old is placed, and the larva is fed with peculiar food, called “royal pap” or “royal jelly,” with the result that in sixteen days—five days less than would be required for a worker and nine less than for a drone—a queen, or perfect female, is produced. She alone has a life extending to years, that of the workers being limited to months at the longest.”
There are many other points to be considered in this matter of honeycomb-construction and the cell-making instinct of the hive-bee, but I will rest content with the points I have already raised, merely asking any unprejudiced person whether Darwin's premises and conclusion are borne out in this one instance by his theory of natural selection. In Brazil there is a bee that builds its comb on the very outmost twigs of lofty trees, as a protection against climbing and marauding enemies. Wherein does this exhibition of intelligence differ much from the New Guinea natives building their huts in lofty trees for similar protective purposes ? To my mind, very little. Yet we are asked to admit that bees live their life and work in sole accordance with a blind principle of natural selection.