Speaking of females laying male eggs alone, I have already expressed my surprise that bees bestow, on those deposited in royal cells, such care and attention as to feed the worms proceeding from them, and, at the period of transformation, to close them up. But I know not, Sir, why I omitted to observe that, after sealing the royal cells, the workers build them up, and sit on them until the last metamorphosis of the included male[I]. The treatment of the royal cells where fertile workers lay the eggs of drones is very different. They begin indeed with bestowing every care on their eggs and worms; they close the cells at a suitable time, but never fail to destroy them three days afterwards.
Having finished these first experiments with success, I had still to discover the cause of the expansion of the sexual organs of fertile workers. M. Riems had not engaged in this interesting problem; and at first I dreaded that I should have no other guide towards its solution than conjecture. Yet from serious reflection, it appeared, that, by connecting the facts contained in this letter, there was some light that might elucidate my procedure in this new research.
From M. Schirach's elegant discoveries, it is beyond all doubt that common bees are originally of the female sex. They have received from nature the germs of an ovary, but she has allowed its expansion only in the particular case of their receiving a certain aliment while a worm. Thus it must be the peculiar object of inquiry whether the fertile workers get that aliment while worms.
All my experiments convince me that bees, capable of laying, are produced in hives that have lost the queen. A great quantity of royal jelly is then prepared for feeding the larvæ destined to replace her. Therefore, if fertile workers are produced in this situation alone, it is evident their origin is only in those hives where bees prepare the royal jelly. Towards this circumstance, I bent all my attention. It induced me to suspect that when bees give the royal treatment to certain worms, they either by accident or a particular instinct, the principle of which is unknown to me, drop some particles of royal jelly into cells contiguous to those containing the worms destined for queens. The larvæ of workers that have accidentally received portions of so active an aliment, must be more or less affected by it; and their ovaries should acquire a degree of expansion. But this expansion will be imperfect; why? because the royal food has been administered only in small portions, and, besides, the larvæ having lived in cells of the smallest dimensions, their parts cannot extend beyond the ordinary proportions. Thus, the bees produced by them will resemble common workers in size and all the external characteristics. Added to that, they will have the faculty of laying some eggs, solely from the effect of the trifling portion of royal jelly mixed with their aliment.
That we may judge of the justness of this explanation, it is necessary to consider fertile workers from their origin; to investigate whether the cells, where they are brought up, are constantly in the vicinity of the royal cells, and if their food is mixed with particles of the royal jelly. Unfortunately, the execution of these experiments is very difficult. When pure, the royal jelly is recognised by its sharp and pungent taste; but, when mixed with other substances, the peculiar savour is very imperfectly distinguished. Thus I conceived, that my investigation should be limited to the situation of the cells; and, as the subject is important, permit me to enter a little into detail[J].
In June 1790, I observed that one of my thinnest hives had wanted the queen several days, and that the bees had no mean of replacing her, there being no workers' worms. I then provided them with a small portion of comb, each cell containing a young worm of the working species. Next day, the bees prolonged several cells around the worms destined for queens, in the form of royal ones. They also bestowed some care on the worms in the adjoining cells. Four days afterwards, all the royal cells were shut, and we counted nineteen small cells also perfected and closed by a covering almost flat. In these were worms that had not received the royal treatment; but as they had lived in the vicinity of the worms destined for replacing the queens, it was very interesting to follow their history, and necessary to watch the moment of their last transformation. I removed the nineteen cells into a grated box, which was introduced among the bees. I also removed the royal cells, for it was of great importance, that the queens they would produce should not disturb or derange the result of the experiment. But here another precaution was also requisite. It was to be feared, that the bees being deprived of the produce of their labour, and the object of their hope might be totally discouraged; therefore, I supplied them with another piece of comb, containing the brood of workers, reserving power to destroy the young brood when necessary. This plan succeeded admirably. The bees, in bestowing all their attention on these last worms, forgot those that had been removed.
When the moment of transformation of the nymphs in the nineteen cells arrived, I examined the grated box frequently every day, and at length found six bees exactly similar to common bees. The worms of the remaining thirteen had perished without changing.
The portion of brood comb that had been put into the hive to prevent the discouragement of the bees was then removed. I put aside the queens produced in the royal cells; and having painted the thorax of the six bees red, and amputated the right antenna, I transferred the whole six into the hive, where they were well received.
You easily conceive my object, Sir, in this course of observations. I knew there was neither a large nor small queen in the hive: therefore, if, in the sequel, I should find new laid eggs in the combs, how very probable must it be that they had been produced by some of the six bees? But, to attain absolute certainty, it was necessary to take them in the act of laying. Some ineffaceable mark was also required for distinguishing them in particular.
This proceeding was attended with the most ample success. We soon found eggs in the hive; their number increased daily; and their worms were all drones. But a long interval elapsed before we could take the bees that laid them. At length, by means of assiduity and perseverance, we perceived one introducing the posterior part into a cell; we opened the hive, and caught the bee: We saw the egg it had deposited, and by the colour of the thorax, and privation of the right antenna, instantly recognised that it was one of the six that had passed to the vermicular state in the vicinity of the royal cells.