The peculiarities of this experiment seem worthy of attention. It proves that nature does not allow the queen the choice of the eggs she is to lay. It is ordained that, at a certain time of the year, she shall produce those of males, and at another time the eggs of workers, and this order cannot be inverted. We have seen that another fact led me to the same consequence; and as that was extremely important, I am delighted to have it confirmed by a new observation. Let me repeat, therefore, that the eggs are not indiscriminately mixed in the ovaries of the queen, but arranged so that, at a particular season, she can lay only a certain kind. Thus, it would be vain at that time of the year, when the queen should lay the eggs of workers, to attempt forcing her to lay male eggs, by filling the hives with large cells; for, by the experiment just described, we learn, that she will rather drop the workers eggs by chance than deposit them in an unsuitable place; and that she will not lay the eggs of males. I cannot yield to the pleasure of allowing this queen discernment or foresight, for I observe a kind of inconsistency in her conduct. If she refused to lay the eggs of workers in large cells, because nature has instructed her that their size is neither proportioned to the size nor necessities of common worms, would not she also have been instructed not to lay several eggs in one cell?
It seems much easier to rear a worker's worm in a large cell, than to rear several of the same species in a small one. Therefore, the supposed discrimination of bees is not very conspicuous. Here the most prominent feature of industry appears in the common bees. When I supplied them with a comb of small cells, full of male brood, their activity was awakened; but instead of bestowing the necessary care on this brood, as they would have done in every other situation, they destroyed the whole nymphs and larvae, and cleaned out their cells, that the queen, now oppressed with the necessity of laying, might suffer no delay in depositing her eggs. Could we allow them either reason or reflection, this would be an interesting proof of their affection for her.
The experiment, now detailed at length, not having fulfilled my object in determining the influence of the size of the cells on that of the worms, I invented another which proved more successful.
Having selected a comb of large cells, containing the eggs and worms of males, I removed all the worms from their farina, and my a.s.sistant subst.i.tuted those of workers a day old in their place. Then he introduced this comb into a hive that had the queen. The bees did not abandon these subst.i.tuted worms; they covered their cells with a top almost flat, a kind quite different from what is put on the cells of males; which proves, that they were well aware that these, though inhabiting large cells, were not males. This comb remained eight days in the hive, counting from the time the cells were sealed. I then removed it to examine the included nymphs, which proved those of workers in different stages of advancement; but, as to size and figure, they perfectly resembled what had grown in the smallest cells. I thence concluded, that the larvae of workers do not acquire greater size in large than in small cells. Although this experiment was made only once, it seems decisive. Nature has appropriated cells of certain dimensions for the worms of workers while in their vermicular state; undoubtedly she has ordained that their organs should be fully expanded, and there is sufficient s.p.a.ce for that purpose; therefore more would be useless.
Their expansion ought to be no greater in the most s.p.a.cious cells than in those appropriated for them. If some cells smaller than common ones are found in combs, and the eggs of workers are deposited there, the size of the bees will probably be less than that of common workers, because they have been cramped in the cells; but it does not thence ensue, that a larger cell will admit of them growing to a greater size.
The effect produced on the size of drones by the size of the cells their worms inhabit, may serve as a rule for what should happen to the larvae of workers in the same circ.u.mstances. The large cells of males are sufficiently capacious for the perfect expansion of their organs. Thus, although reared in cells of still greater capacity, they will grow no larger than common drones. We have had evidence of this in those produced by queens whose fecundation has been r.e.t.a.r.ded. You will remember, Sir, that they sometimes lay male eggs in the royal cells.
Now, the males proceeding from them, and reared in cells much more s.p.a.cious than nature has appropriated for them, are no larger than common males. Therefore it is certain, that whatever be the size of the cells where the worms acquire their increment, the bees will attain no greater size than is peculiar to their species. But if, in their primary form, they live in cells smaller than they should be, as their growth will be checked, they will not attain the usual size, of which there is proof in the following experiment. I had a comb consisting of the cell of large drones, and one with those of workers, which also served for the male worms. Of these, my a.s.sistant took a certain number from the smallest cells, and deposited them on a quant.i.ty of food purposely prepared in the large ones; and in return he introduced into the small cells the worms that had been hatched in the other, and then committed both to the care of the workers in a hive where the queen laid the eggs of males only. The bees were not affected by this change; they took equal care of the worms; and when the period of metamorphosis arrived, gave both kinds that convex covering usually put on those of the males.
Eight days afterwards, we removed the combs, and found, as I had expected, nymphs of large males in the large cells, and those of small males in the small ones.
You suggested another experiment which I carefully made, but it met with an unforeseen obstacle. To appreciate the influence of the royal food on the expansion of the worms, you desired me to supply the worm of a worker in a common cell with it. Twice I have attempted this operation without success. Nor do I think it can ever succeed. If bees get the charge of worms, in whose cells the royal food is deposited, and if at the same time they have a queen, they soon remove the worms and greedily devour the food. When, on the contrary, they are deprived of a queen, they change the cells containing worms into cells of the largest kind.
Then the worms will infallibly be converted to queens.
But there is another situation where we can judge of the influence of the royal food administered to worms in common cells. I have spoken at great length in my letter on the existence of fertile workers. You cannot forget, Sir, that the expansion of their s.e.xual organs is owing to the reception of some particles of royal jelly, while in the vermicular form. For want of new observations, I must refer you to what is previously said on the subject.
_PREGNY, 4 September 1791._
LETTER IX.
_ON THE FORMATION OF SWARMS._
I can add but a few facts to the information M. de Reaumur has communicated relative to swarms.
A young queen, according to this celebrated naturalist, is always or almost always at the head of a swarm; but he does not a.s.sert the fact positively, and had some doubts on the subject. "Is it certain," says he "as we have hitherto supposed, in coincidence with all who have treated of bees, that the new colony is always conducted by a young mother? May not the old mother be disgusted with her habitation? or may she not be influenced by some particular circ.u.mstances to abandon all her possessions to the young female? I wish it had been in my power to solve this question otherwise than by mere probabilities, and that some misfortune had not befallen all the bees whose queen I had marked red on the thorax."
These expressions seem to indicate, that M. de Reaumur suspected that the old queens sometimes conducted the young swarms. By the following details, you will observe, that his suspicions are fully justified.
In the course of spring and summer, the same hive may throw several swarms. The old queen is always at the head of the first colony; the others are conducted by young queens. Such is the fact which I shall now prove; and the peculiarities attending it shall be related.
But previous to entering on this subject, I should repeat what has already been frequently observed, that the _leaf_ or flat hives are indispensible in studying the industry and instinct of bees. When they are left at liberty to conduct several rows of parallel combs, we can no longer observe what is continually pa.s.sing between them, or they must be dislodged by water or smoke, for examining what has been constructed; a violent proceeding, which has a material influence on their instinct, and consequently exposes an observer to the risk of supposing simple accidents permanent laws.
I now proceed to experiments proving that an old queen always conducts the first swarm.
One of my gla.s.s hives consisted of three parallel combs, placed in squares that opened like the leaves of a book. It was well peopled and abundantly provided with honey, wax, and brood, of every age. On the fifth of May 1788, I removed the queen, and on the sixth, transferred all the bees into another hive, with a fertile queen at least a year old. They entered easily and without fighting, and were in general well received. The old inhabitants of the hive, which, since privation of their queen, had begun twelve royal cells, also gave the fertile queen a good reception; they presented her with honey, and formed regular circles around her. However, there was a little agitation in the evening, but confined to the surface of the comb where we had put the queen, and which she had not quitted. All was perfectly quiet on the other side of this comb.
In the morning of the seventh, the bees had destroyed the twelve royal cells, but, independent of that, order continued prevalent in the hive; the queen laid the eggs of males in the large cells, and those of workers in the small ones, respectively.
Towards the twelfth, we found the bees occupied in constructing twenty-two royal cells, of the same species described by M. de Reaumur, that is the bases not in the plane of the comb, but appended perpendicularly by pedicles or stalks of different length, like stalact.i.tes, on the edge of the pa.s.sage made by the bees through their combs. They bore considerable resemblance to the cup of an acorn, and the longest were only about two lines and a half in depth from the bottom to the orifice.
On the thirteenth, the queen seemed already more slender than when introduced into the hive; however she still laid some eggs, both in common cells and those of males. We also surprised her this day laying in a royal cell: she first dislodged the worker there employed, by pushing it away with her head, and then supported herself by the adjoining cells while depositing the egg.
On the fifteenth, the queen was still more slender: the bees continued their attention to the royal cells, which were all unequally advanced; some to three or four lines in height, while others were already an inch long; which proved that the queen had not laid in the whole at the same time.
At the moment when least expected, the hive swarmed on the nineteenth; we were warned of it by the noise in the air; and hastened to collect and put the bees into a hive purposely prepared. Though we had overlooked the facts attending the departure of the swarm, the object of this experiment was fulfilled; for, on examination of all the bees, we were convinced they had been conducted by the old queen; by that we introduced on the sixth of the month, and which had been deprived of one of the antennae. Observe, there was no other queen in this colony. In the hive she had left, we found seven royal cells close at the top, but open at the side, and quite empty. Eleven more were sealed; and some others newly begun; no queen remained in the hive.
The new swarm next became the object of our attention: we observed it during the rest of the year, during winter and the subsequent spring; and, in April, we had the satisfaction of seeing a new swarm depart with the same queen at its head that had conducted the former swarm in May the preceding year.
You will remark, Sir, that this experiment is positive. We put an old queen in a gla.s.s hive while laying the eggs of males. The bees received her well, and at that time began to construct royal cells; she laid in one of them before us; and in the last place led forth the swarm.
We have several times repeated the same experiment with equal success.
Thus it appears incontestible, that the old queen always conducts the first swarm; but never quits the hive before depositing eggs in the royal cells, from which other queens will proceed after her departure.
The bees prepare these cells only while the queen lays male eggs; and a remarkable fact attends it, that after this laying terminates, her belly being considerably diminished, she can easily fly, whereas, her belly is previously so heavy she can hardly drag it along. Therefore it is necessary she should lay in order to be in a condition for undertaking her journey, which may sometimes be very long.
But this single condition is not enough. It is also requisite that the bees be very numerous: they should even be superabundant, and a person might say they are aware of it: for, if the hive is thin, no royal cells are constructed when the male eggs are laid, which is solely at the period that the queen is able to conduct a colony. This fact was proved by the following experiment on a large scale.
On the third of May 1788, we divided eighteen hives into two portions; all the queens were about a year old. Thus each portion of the hives had but half the bees that were originally there. Eighteen halves wanted queens, but the other eighteen had very fertile ones. They soon began to lay the eggs of males; but, the bees being few, they did not construct royal cells, and none of the hives threw a swarm.--Therefore, if the hive containing the old queen is not very populous, she remains in it until the subsequent spring; and if the population is then sufficient, royal cells will be constructed: the queen will begin to lay male eggs, and, after depositing them, will issue forth at the head of a colony, before the young queens are produced.
Such is a very brief abstract of my observations on swarms conducted by old queens. You must excuse the long detail on which I am about to enter, concerning the history of the royal cells left by the queen in the hive. Every thing relative to this part of the history of bees has. .h.i.therto been very obscure. A long course of observations, protracted even during several years, was necessary to remove, in some degree, the veil that concealed these mysteries. I have been indemnified for the trouble, indeed, by the pleasure of seeing my experiments reciprocally confirmed; but, considering the a.s.siduity required in these researches, they were truly very laborious.
Having established in 1788 and 1789, that queens a year old conducted the first swarm, and that they left worms or nymphs in the hive to transform into queens in their turn; I endeavoured, in 1790, to profit by the goodness of the spring, to study all that related to these young queens; and I shall now extract the chief experiments from my journal.
On the fourteenth of May, we introduced two portions of bees, from the straw hives, into a large gla.s.s hive very flat; and allowed them only one queen of the preceding year, and which had already commenced laying in its native hive. We introduced her on the fifteenth. She was very fertile. The bees received her well, and she soon began to lay in large and small cells alternately.
On the twentieth, we saw the formation of twelve royal cells, all on the edges of the communications, or pa.s.sages through the combs, and shaped liked stalact.i.tes.
On the twenty-seventh, ten were much but unequally enlarged; but none so long as when the worms are hatched.
On the twenty-eighth, previous to which the queen had not ceased laying, her belly was very slender, and she began to exhibit signs of agitation.
Her motion soon became more lively, yet she still continued examining the cells as when about to lay; sometimes introducing half her belly, but suddenly withdrawing it, without having laid. At other times she deposited an egg, which lay in an irregular position, on one side of the hexagon, and not fixed by an end to the bottom of the cell. The queen produced no distinct sound in her course, and we heard nothing different from the ordinary humming of bees. She pa.s.sed over those in her way; sometimes when she stopped, the bees meeting her also stopped; and seemed to consider her. They advanced briskly, struck her with their antennae, and mounted on her back. She then went on carrying some of the workers on her back. None gave her honey, but she voluntarily took it from the cells in her way. The bees no longer inclosed and formed regular circles around her. The first, aroused by her motions, followed her running in the same manner, and in their pa.s.sage excited those still tranquil on the combs. The way the queen had traversed was evident after she left it, by the agitation created, which was never afterwards quelled: she had soon visited every part of the hive, and occasioned a general agitation; if some places still remained tranquil, the bees in agitation arrived, and communicated their motion. The queen no longer deposited her eggs in cells; she let them fall fortuitously: nor did the bees any longer watch over the young; they ran about in every different direction; even those returning from the fields, before the agitation came to its height, no sooner entered the hive than they partic.i.p.ated in these tumultuous motions. They neglected to free themselves of the waxen pellets on their limbs, and ran blindly about. At last the whole rushed precipitately towards the outlets of the hive, and the queen along with them.
As it was of much consequence to see the formation of new swarms in this hive, and, for that reason, as I wished it to continue very populous, I removed the queen, at the moment she came out, that the bees might not fly too far, and that they might return. In fact, after losing their female, they did return to the hive. To increase the population still more, I added another swarm, which had come from a straw hive on the same morning, and removed its queen also.
All these facts were certain, and appeared susceptible of no error.
Notwithstanding this, I was particularly earnest to learn whether old queens always followed the same course; which induced me, on the twenty-ninth, to replace, in the gla.s.s hive, the queen a year old, which had hitherto been the subject of my experiments, and had just began to lay the eggs of males. On the same day, we found one of the royal cells left by the preceding queen larger than the rest; and, from its length, supposed the included worm two days old: the egg had, therefore, been laid on the twenty-fourth by that queen, and the worm was hatched on the twenty-seventh. On the thirtieth, the queen laid a great deal in the large and small cells alternately. Now, and the two following days, the bees enlarged several royal cells, but unequally, which proved that they included larvae of different ages. One was closed on the first of June, and on the second another. The bees also commenced some new ones. All was perfectly quiet at eleven in the morning; but, at mid-day, the queen, from the utmost tranquillity, became evidently agitated; and her agitation insensibly communicated to the workers in every part of their dwelling. In a few minutes they precipitately crowded to the entrances, and, along with the queen, left the hive. After they had settled on the branch of a neighbouring tree, I sought for the queen; thinking that, by removing her, the bees might return to the hive, which actually ensued.
Their first care seemed to consist in seeking the female; they were still in great agitation, but gradually calmed; and in three hours complete tranquillity was restored.
They had resumed their usual occupations on the third: they attended to the young, worked within the open royal cells, and also watched on those that were shut. They made a waved work on them, not by applying wax cordons, but by removing wax from the surface. Towards the top this waved work is almost imperceptible; it becomes deeper above, and the workers excavate it still more from thence to the base of the pyramid.
The cell, when once shut, also becomes thinner; and is so much so, immediately preceding the queen's metamorphosis from a nymph, that all its motions are perceptible through the thin covering of wax on which the waved work is founded. It is a very remarkable circ.u.mstance, that in making the cells thinner, from the moment they are closed, the bees know to regulate their labour so that it terminates only when the nymph is ready to undergo its last metamorphosis.
On the seventh day the cocc.o.o.n is almost completely _unwaxed_, if I may use the expression, at the part next to the head and thorax of the queen. This operation facilitates her exit; for she has nothing to do but cut the silk that forms the cocc.o.o.n. Most probably the object is, to promote evaporation of the superabundant fluids of the nymph. I have made some direct experiments to ascertain the fact, but they are yet unfinished. A third royal cell was closed by the bees on the same day, the third of June, twenty-four hours after closing the second. The like was done to other royal cells successively, during the subsequent days.
Every moment of the seventh, we expected the queen to leave the royal cell shut on the thirtieth of May. The seven days had elapsed. The waving of her cell was so deep, that what pa.s.sed within was pretty perceptible; we could discern that the silk of the cocc.o.o.n was cut circularly, a line and a half from the extremity; but the bees being unwilling that she should yet quit her cell, they had soldered the covering to it with some particles of wax. What seemed most singular was, that this female emitted a very distinct sound, or clacking from her prison. It was still more audible in the evening, and even consisted of several monotonous notes in rapid succession.
The same sound proceeded from the royal cell on the eighth. Several bees kept guard round each royal cell.
The first cell opened on the ninth. The young queen was lively, slender, and of a brown colour. Now, we understood why bees retain the female captive in their cells, after the period for transformation has elapsed; it is, that they may be able to fly the instant they are hatched. The new queen occupied all our attention. When she approached the other royal cells, the bees on guard pulled, bit her, and chased her away; they seemed to be greatly irritated against her, and she enjoyed tranquillity only when at a good distance from these cells. This procedure was frequently repeated through the day. She twice emitted the sound; in doing so she stood, her thorax against a comb, and the wings crossed on her back; they were in motion but without being unfolded or further opened. Whatever might be the cause of her a.s.suming this att.i.tude, the bees were affected by it; all hung down their heads, and remained motionless.
The hive presented the same appearances on the following day.
Twenty-three royal cells yet remained, a.s.siduously guarded by a great many bees. When the queen approached, all the guards became agitated, surrounded her on all sides, bit, and commonly drove her away; sometimes when in these circ.u.mstances, she emitted her sound, a.s.suming the position just described, from that moment the bees became motionless.
The queen confined in the second cell had not yet left it, and was heard to hum several times. We accidentally discovered how the bees fed her.
On attentive examination, a small aperture was perceptible in the end of the cocc.o.o.n which she had cut to escape, and which her guards had again covered with wax, to confine her still longer. She thrust her trunk through the cleft; at first the bees did not observe it alternately thrust out and drawn in, but one at length perceiving it, came to apply its trunk to that of the captive queen, and then gave way to others that also approached her with honey. When satisfied she retracted her trunk, and the bees again closed up the opening with wax.