In relating my first observations on queens that lay male eggs alone, I have proved that they lay them in cells of all dimensions indifferently, and even in royal cells. It is also proved that the same treatment is given to male worms hatched from eggs laid in the royal cells, as if they were actually to be transformed to queens; and I have added, that in this instance the instinct of the workers appeared defective. It is indeed most singular, that bees which know the worms of males so well when the eggs are laid in small cells, and never fail to give them a convex covering when about to transform to nymphs, should no longer recognise the same species of worms when the eggs are laid in royal cells, and treat them exactly as if they should change to queens. This irregularity depends on something I cannot comprehend.
In revising what is said on this subject, I observe still wanting an interesting experiment to complete the history of queens that lay only the eggs of drones. I had to investigate whether these females could themselves distinguish that the eggs they deposit in the royal cells would not produce queens. I have already observed that they do not endeavour to destroy these cells when close, and I thence concluded, that in general the presence of royal cells in their hive does not inspire them with the same aversion to females whose fecundation has been retarded; but to ascertain the fact more correctly, it was essential to examine how the presence of a cell containing a royal nymph would affect a queen that had never laid any other than the eggs of drones.
This experiment was easy; and I put it in practice on the fourth of September, in a hive some time deprived of its queen. The bees had not failed to construct several royal cells for replacing their females. I chose this opportunity for supplying them with a queen, whose fecundation had been retarded to the twenty-eighth day, and which laid none but the eggs of males. At the same time, I removed all the royal cells, except one that had been sealed five days. One remaining was enough to shew the impression it would make on the stranger queen introduced; had she endeavoured to destroy it; this, in my opinion, would have proved that she anticipated the origin of a dangerous rival. You must admit the use I make of the word anticipate; it saves a long paraphrase; I feel the impropriety of it. If, on the contrary, she did not attack the cell I would thence conclude that the delay of fecundation, which deprived her of the power of laying workers eggs, had also impaired her instinct. This was the fact; the queen passed several times over the royal cell, both the first and the subsequent day, without seeming to distinguish it from the rest. She quietly laid in the surrounding cells; notwithstanding the cares incessantly bestowed by the bees upon it, she never one moment appeared to suspect the danger with which the included royal nymph threatened her. Besides, the workers treated their new queen as well as they would have treated any other female. They were lavish of honey and respect, and formed those regular circles around her that seem an expression of homage.
Thus, independent of the derangement occasioned by retarded impregnation, in the sexual organs of queens, it certainly impairs their instinct. Aversion or jealousy is no longer preserved against their own sex in the nymphine state, nor do they longer endeavour to destroy them in their cradles.
My readers will be surprised that queens whose fecundation has been retarded, and whose fecundity is so useless to bees, should be so well treated and become as dear to them as females laying both kinds of eggs. But I remember to have observed a fact more astonishing still. I have seen workers bestow every attention on a queen though sterile; and after her death treat her dead body as they had treated herself when alive, and long prefer this inanimate body to the most fertile queens I had offered them. This sentiment, which assumes the appearance of so lively an affection, is probably the effect of some agreeable sensation communicated to bees by their queen, independent of fertility. Those laying only the eggs of males probably excite the same sensation in the workers.
I now recollect that the celebrated Swammerdam somewhere observes, that when a queen is blind, sterile, or mutilated, she ceases to lay, and the workers of her hive no longer labour or make any collections, as if aware that it was now useless to work. He cites no experiment that led him to the discovery. Those made by myself have afforded some very singular results.
I frequently amputated the four wings of queens; and not only did they continue laying, but the same confederation of them was testified by the workers as before. Therefore, Swammerdam has no foundation for asserting, that mutilated queens cease to lay. Indeed, from his ignorance of fecundation taking place without the hives, it is possible he cut the wings off virgin queens, and they, becoming incapable of flight, remained sterile from inability to seek the males in the air. Thus, amputation of the wings does not produce sterility in queens.
I have frequently cut off one [antennæ] to recognise a queen the more easily, and it was not prejudicial to her either in fecundity or instinct nor did it affect the attention paid to her by the bees. It is true, that as one still remained, the mutilation was imperfect; and the experiment decided nothing. But amputation of both antennæ produced most singular effects. On the fifth of September, I cut both off a queen that laid the eggs of males only, and put her into the hive immediately after the operation. From this moment there was a great alteration in her conduct. She traversed the combs with extraordinary vivacity. Scarcely had the workers time to separate and recede before her; she dropped her eggs, without attending to deposit them in any cell. The hive not being very populous, part was without comb. Hither she seemed particularly earnest to repair, and long remained motionless. She appeared to avoid the bees; however, several workers followed her into this solitude, and treated her with the most evident respect. She seldom required honey from them, but, when that occurred, directed her trunk with an uncertain kind of feeling, sometimes on the head and sometimes on the limbs of the workers, and if it did reach their mouths, it was by chance. At other times she returned upon the combs, then quitted them to traverse the glass sides of the hive: and always dropped eggs during her various motions. Sometimes she appeared tormented with the desire of leaving her habitation. She rushed towards the opening, and entered the glass tube adapted there; but the external orific being too small, after fruitless exertion, she returned. Notwithstanding these symptoms of delirium, the bees did not cease to render her the same attention as they ever pay to their queens, but this one received it with indifference. All that I describe appeared to me the consequence of amputating the antennæ. However, her organization having already suffered from retarded fecundation, and as I had observed her instinct in some degree impaired, both causes might possibly concur in producing the same effect. To distinguish properly what belonged to the privation of the antennæ, a repetition of the experiment was necessary, in a queen otherwise well organised, and capable of laying both kinds of eggs.
This I did on the sixth of September. I amputated both the antennæ of a female which had been several months the subject of observation, and being of great fecundity had already laid a considerable number of workers eggs, and those of males. I put her into the same hive where the queen of the preceding experiment still remained, and she exhibited precisely the same marks of delirium and agitation, which I think it needless to repeat. I shall only add, that to judge better of the effect produced by privation of the antennæ, on the industry and instinct of bees, I attentively considered the manner in which these two mutilated queens treated each other. You cannot have forgot, Sir, the animosity with which queens, possessing all their organs, combat, on which account it became extremely interesting to learn whether they would experience the same reciprocal aversion after losing their antennæ. We studied these queens a long time; they met several times in their courses, and without exhibiting the smallest resentment. This last instance is, in my opinion, the most complete evidence of a change operated in their instinct.