Hitherto no other effect has been observed from the retarded impregnation of animals, but that of rendering them absolutely sterile. The first instance of a female still preserving the faculty of engendering males, is presented by the queen bee. But as no fact in nature is unique, it is most probable that the same peculiarity will also be found in other animals. An extremely curious object of research would be to consider insects in this new point of view, I say insects, for I do not conceive that any thing analogous will be found in other species of animals. The experiments now suggested would necessarily begin with insects the most analogous to bees; as wasps, humble bees, mason bees, all species of flies, and the like. Some experiments might also be made on butterflies; and, perhaps, an animal might be found whose retarded fecundation would be attended with the same effects as that of queen bees. Should the animal be larger, dissection will be more easily accomplished; and we may discover what happens to the eggs when retarded fecundation prevents their expansion. At least, we might hope that some fortunate circumstance would lead to solution of the problem[G].
Let us now return to my experiments. In May 1789, I took two queens just when they had undergone the last metamorphosis: one was put in a leaf hive, well provided with honey and wax, and sufficiently inhabited by workers and males. The other was put into a hive exactly similar, from which all the drones were removed. The entrances of these hives were too confined for the passage of the females and drones, but the common bees enjoyed perfect liberty. The queens were imprisoned thirty days; and being then set at liberty, they departed, and returned impregnated. Visiting the hives in the beginning of July, I found much brood, but wholly consisting of the worms and nymphs of males. There actually was not a single worker's worm or nymph. Both queens laid uninterruptedly until autumn, and constantly the eggs of drones. Their laying ended in the first week of November, as that of my other queens.
I was very earnest to learn what would become of them in the subsequent spring, whether they would resume laying, or if new fecundation would be necessary; and if they did lay, of what species the eggs would be. However, the hives being very weak, I dreaded they might perish during winter. Fortunately, we were able to preserve them; and from April 1790, they recommenced laying. The precautions we had taken prevented them from receiving any new approaches of the male. Their eggs were still those of males.
It would have been extremely interesting to have followed the history of these two females still farther, but, to my great regret, the workers abandoned their hives on the fourth of May, and that same day I found both queens dead. No weevils were in the hive, which could disturb the bees; and the honey was still very plentiful: but as no workers had been been produced in the course of the preceding year, and winter had destroyed many, they were too few in spring to engage in their wonted labours, and, from discouragement, deserted their habitation to occupy the neighbouring hives.
In my Journal, I find a detail of many experiments on the retarded impregnation of queen bees, so many, that transcribing the whole would be tedious. I may repeat, however, that there was not the least variation in the principle, and that whenever the copulation of queens was postponed beyond the twenty-first day, the eggs of males only were produced. Therefore, I shall limit my narrative to those experiments that have taught me some remarkable facts.
A queen being hatched on the fourth of October 1789, we put her into a leaf-hive. Though the season was well advanced, a considerable number of males was still in the hive; and it here became important to learn, whether, at this period of the year, they could equally effect fecundation; also, in case it succeeded, whether a laying, begun in the middle of autumn, would be interrupted or continued during winter. Thus, we allowed the queen to leave the hive. She departed, indeed, but made four and twenty fruitless attempts before returning with the evidence of fecundation. Finally, on the thirty-first of October, she was more fortunate: She departed, and returned with the most undoubted proof of the success of her amours: She was now twenty-seven days old, consequently fecundation had been retarded. She ought to have begun laying within forty-six hours, but the weather was cold, and she did not lay; which proves, as we may cursorily remark, that refrigeration of the atmosphere is the principal agent that suspends the laying of queens during winter. I was excessively impatient to learn whether, on the return of spring, she would prove fertile, without a new copulation. The means of ascertaining the fact was easy; for the entrances of the hives only required contraction, so as to prevent her from escaping. She was confined from the end of October until May. In the middle of March, we visited the combs, and found a considerable number of eggs, but, none being yet hatched, we could not know whether they would produce workers or males. On the fourth of April, having again examined the state of the hive, we found a prodigious quantity of nymphs and worms, all of drones; nor had this queen laid a single worker's egg.
Here, as well as in the preceding experiment, retardation had rendered the queens incapable of laying the eggs of workers. But this result is the more remarkable, as the queen did not commence laying until four months and a half after fecundation. It is not rigorously true, therefore, that the term of forty-six hours elapses between the copulation of the female and her laying; the interval may be much longer, if the weather grows cold. Lastly, it follows, that although cold will retard the laying of a queen impregnated in autumn, she will begin to lay in spring without requiring new copulation.
It may be added, that the fecundity of the queen, whose history is given here, was astonishing. On the first of May, we found in her hive, besides six hundred males, already flies, two thousand four hundred and thirty-eight cells, containing either eggs or nymphs of drones. Thus, she had laid more than three thousand male eggs during March and April, which is above fifty each day. Her death soon afterwards unfortunately interrupted my observation, I intended to calculate the total number of male eggs that she should lay throughout the year, and compare it with those of queens whose fecundation had not been retarded. You know, Sir, that the latter lay about two thousand male eggs in spring; and another laying, but less considerable, commences in August, also in the interval, that they produce the eggs of workers almost solely. But it is otherwise with the females whose copulation has been retarded: they produce no workers' eggs. For four or five months following, they lay the eggs of males without interruption, and in such numbers, that, in this short time, I suppose one queen gives birth to more drones than a female, whose fecundation has not been retarded, produces in the course of two years. It gives me much regret, that I have not been able to verify this conjecture.
I should also describe the very remarkable manner in which queens, that lay only the eggs of drones, sometimes deposit them in the cells. Instead of being placed in the lozenges forming the bottom, they are frequently deposited on the lower side of the cells, two lines from the mouth. This arises from the body of such queens being shorter than that of those whose fecundation has not been retarded. The extremity remains slender, while the first two rings next the thorax are uncommonly swoln. Thus, in disposing themselves for laying, the extremity cannot reach the bottom of the cells on account of the swoln rings; consequently the eggs must remain attached to the part that the extremity reaches. The worms proceeding from them pass their vermicular state in the same place where the eggs were deposited, which proves that bees are not charged with the care of transporting the eggs as has been supposed. But here they follow another plan. They extend beyond the surface of the comb those cells where they observe the eggs deposited, two lines from the mouth.
Permit me, Sir, to digress a moment from the subject, to give the result of an experiment which seems interesting. Bees, I say, are not charged with the care of transporting into cells, the eggs misplaced by the queen: and, judging by the single instance I have related, you will think me well entitled to deny this feature of their industry. However, as several authors have maintained the reverse, and even demanded our admiration of them in conveying the eggs, I should explain clearly that they are deceived.