BEE-spoke-data/bees-v0 huggingface dataset:
8 The experiments which I suggest in this paragraph remind me of a singular reflection of Mr. de Reaumur. Speaking of viviparous flies, he says it might not be impossible for a hen to give life to a living chick, if after fecundation, the eggs that she might lay were retained for 20 days in her oviducts. (See Reaumur sur les Insects, tome IV, M&moire 10.)I feared that they might perish during the winter. Fortunately, we succeeded in preserving them, and, as early as April, they recommended their laying: through the precautions we had taken we were sure that they had not received new approaches of the male; their eggs were still drone eggs.
It would have been very interesting to follow farther the history of these females, but, to my great regret, their workers abandoned them on the 4th of May, and on that same day we found both the queens dead. Yet there were no moth worms in the combs to cause any disturbance to the bees, and the honey was still fairly abundant, but, as in the course of the previous year no workers had been born, and as winter had destroyed many, they found themselves in too small numbers to engage in their ordinary labors, and in their discouragement they deserted their hives to join the neighboring hives.
I find, in my journal, the detail of a multitude of experiments on the retarded foundation of queen bees: it would be an endless task to transcribe them all here: I repeat that there was not the least variation in the principal result and every time and whenever the mating of the queens was delayed beyond the 21st day, they laid none but male eggs. Therefore, sir, I will limit myself to expound to you those of my experiments which have taught me some remarkable facts not yet mentioned.
On the 4th of October 1789, a queen hatched in one of my hives: we placed her in a leaf hive. Although the season was well advanced there were still many males in the hives. It was important to learn whether, at this time of the year, they could equally effect decundation, and, in case of success, whether the laying, begun in the middle of autumn, would be interrupted or continued during the winter. So we allowed this queen to leave the hive. She did, but made 24 fruitless trips before returning with the signs of decundation. At last, on the 31st of October, she was more fortunate, she came back with most evident marks of the success of her amours; she was then 27 days old, so that her decundation had been much delayed. She should have begun laying 46 hours afterwards, but the weather was cold and she did not lay; which, let me say it in passing, proves that refrigeration of the temperature is the principal cause of the suspension of the queens' laying in autumn. I was very impatient to learn whether, on the return of spring, she would prove fertile, without the need of a new union. The means of ascertaining this was easy; for it only required contraction of the entrance of her hive, so that she might not escape. I confined her from the end of October until May. In the middle of March we examined her combs, and found many eggs, but as they were laid in cells of the small size, we had to wait a few days to pass judgment. On the 4th of April, we again examined the hive and found a prodigious quantity of nymphs and worms. They were all of drones; the queen had not laid a single worker egg.
Here, as in former experiments, delayed fecundation had rendered the queen bee incapable of laying eggs of workers. This result is the more remarkable that the laying of the queen began four months and a half after fecundation. The term of 46 hours which usually elapses between mating and laying is therefore not a rigorous term; the interval may be much
diaphragm, which left on each side a sufficient space for the passage of the workers from the one story to the other, but too narrow for the queen. I stocked this hive with a good number of bees, and confined in the upper story a very prolific female that had just lately finished her great laying of male eggs. Therefore, this female had only worker eggs to lay; she was compelled to deposit them in large cells, since there were none of the other sort about her. You will divine, sir, the aim which I proposed in thus disposing matters. My reasoning was very simple. If the queen lays worker eggs in large cells and if the workers are charged with transporting them when misplaced, they would not fail to take advantage of the liberty allowed them to pass from one story to the other; they would seek the eggs deposited in large cells and carry them to the lower story, containing the cells suitable for them. If, on the contrary, they left the worker eggs in the large cells, I should obtain certain proof that they are not entrusted with transporting them.
The result of this experiment excited my curiosity deeply. We watched the queen of this hive and her bees, for several days in succession, with sustained attention. During the first 24 hours, the queen obstinately refused to lay a single egg in the large cells which surrounded her; she examined them one after another, but passed on without insinuating her belly within any of them: she seemed restless, distressed; she traversed the combs in all directions; the weight of her eggs appeared to burden her, but she persisted in withholding them rather than deposit them in cells of unsuitable diameter. Her bees, however, did not cease to pay her homage and treat her as a mother. I even saw, with pleasure, that when she approached the edges of the partition which separated the two stories, she gnawed them to try to enlarge the passage; the workers also labored with their teeth, and made efforts to enlarge the entrance to her prison, but ineffectually. On the second day, the queen could no longer retain her eggs, they escaped in spite of her, she dropped them at random. Yet, we found 8 or 10 of them in the cells, but they had disappeared the following day. We then conceived that the bees had transported them into the lower cells, we sought them with the greatest care; but I can assure you that there was not a single one there. On the 3d day the queen again laid a few eggs, which disappeared as the first. We again sought them in the small cells but they were not there. The fact is that the workers eat them, and that is what deceived the former observers who claim that they transport them. They noted the disappearance of the misplaced eggs from the cells, and without farther investigation asserted that the bees place them elsewhere; they take them indeed, but not to transfer them; they eat them.
Thus, nature has not entrusted the bees with the care of placing the eggs in appropriate cells; but she has given the females themselves enough instinct to know the nature of the egg which they are about to lay and to place it in a suitable cell. Mr. de Reaumur had already observed this, and in this my observations agree with his. It is therefore certain that, in the natural state, when feoundation has been timely and the queen has not suffered from any circumstance, she does not err in the choice of the different cells in which she is to deposit her eggs: she does not fail to lay those of workers in the small cells and those of males in the large cells.
I am speaking here, Sir, of what happens in natural conditions.--This distinction is important, for we do not find the same unerring instinct, in the behavior of females whose mating has been too long delayed; they make no selection of cells in which they are to lay. This fact is so true, that when I first observed the queens whose freundation had been delayed, I often made mistakes on the kinds of eggs that they were laying; when I saw them lay indistinctively in both small and large cells, I thought that the eggs laid in the small cells were worker eggs: I was thus very much surprised, at the time of their transformation into nymphs, to see them close up the cells with convex cappings, exactly similar to those that they place upon the cells of males: they were, in fact, males; those that were born in small cells were drones of the small size; those that were born in large cells became large drones.--I warn those who wish to repeat my experiments upon queens laying only male eggs, that they must expect to see those queens deposit drone eggs in worker cells.
Moreover, here is a curious observation: these same females, whose freundation has been delayed, sometimes lay drone eggs in royal cells. When I give the history of swarms, I shall show, that, in a natural condition, when the queens begin their great laying of male eggs, the workers construct numerous royal cells: there is probably a secret relation between the production of those eggs and the formation of those royal cells: it is a law of nature from which the bees never derogate. It is not surprising, therefore, that cells of this sort are constructed in hives governed by queens laying drone eggs only. Neither is it singular that these queens should deposit in royal cells the eggs of the only kind that they are able to lay, for in general their instinct seems altered. But what I cannot comprehend is that the bees take the same care of male eggs deposited in royal cells, as of the eggs that would become queens; they give them a more abundant food, they build up the cells as if containing a royal worm; they work upon them, in fact, with such regularity that we have been frequently deceived, ourselves. More than once we opened one of these cells, after they had sealed them, in the persuasion of finding royal nymphs, but it was always a drone nymph that was lodged there. Here the instinct of the workers appear defective. In the natural state, they distinguish the worms of males from those of common bees, since they never fail to give a particular cover to the drone cells. Why then do they no longer distinguish the drone worms, when they are placed in royal cells? This fact deserves much attention. I am convinced that, in order to understand the laws of the instinct of animals, we must carefully observe the cases where this instinct appears to err. (See note 1 of the 12th letter.)
Perhaps in beginning this letter, Sir, I should have given an abstract of the observations which other naturalists have made previously, on queens laying only male eggs; but I shall here repair this omission.--In a work entitled, Histoire de la reine des abeilles, translated from the German by Blassiere, there is printed a letter which Mr. Sehirach wrote you in date of April 15, 1771, in which he speaks of several of his hives, the brood of which changed into drones. You remember, sir, that he ascribed this accident to some unknown vice in the ovaries of the queen in these hives; but he was far from suspecting that retarded decundation caused this vice of
Nor is this all: bees can convert into queens worms still younger. The following experiment taught me that, when they have lost their queen, they destine worms only a few hours old to replace her. A hive in my possession, deprived of female, had not had any eggs or worms for a long time: I gave them a queen of the greatest fecundity; she soon began to lay in the worker cells; I kept her in that hive a little less than 3 days, and removed her before any of her eggs were hatched: the following day, that is the fourth day, Burnens counted 50 small worms, the oldest of which were hardly 24 hours old. However, several of them were already destined for queens, which was proved by the bees depositing around them a much greater provision of jelly than is supplied to common worms. The following day, the worms being nearly 40 hours old, the bees had enlarged their eradles; they had converted their hexagonal cells into cylindrical cells of the greatest capacity; they still labored at them the following days and closed them on the fifth day from the hatching of the worms. Seven days after the sealing of the first of these royal cells, we saw a queen of the largest size emerge from it. This queen rushed towards the other royal cells and endeavored to destroy the nymphs or worms that were in them. I shall narrate the effects of her fury, in another letter.
You see, Sir, from these details, that Mr. Schirach had not yet sufficiently varied his experiments, when he asserted that, in order to become converted into queens, the worms of workers should be 3 days old. It is undoubted that the operation is equally successful, not only with worms 2 days old, but even with those that are only a few hours old.
After having made the researches which I have just described, to verify Mr. Schirach's discovery, I desired to ascertain whether, as this observer claims, the only means which the bees have of procuring a queen is to give the common worms a certain kind of food and rear them in larger cells. You have not forgotten that Mr. de Reaumur had very different ideas upon this. "The mother," said he, "must lay and does lay eggs from which bees fit to become mothers must in their turn proceed. She does so and we will show that the workers know what she is to do. The bees, to which mothersare so precious, seem to take great interest in the eggs which are to supply them, and consider them as very important: they construct particular cells in which they are to be deposited, &c. When a royal cell is only just begun, it has the shape of a cup, or more precisely of an acorn cup, from which the acorn has dropped, &c, &c."
Mr. de Reaumur did not suspect the possibility of the conversion of a common worm into a queen, but he conceived that the mother bee laid a particular species of eggs in the royal cells, from which came worms that should be queens in their turn. On the contrary, according to Mr. Schirach, bees always having the possibility of procuring a queen, by rearing worker worms 3 days old in a particular manner, it would be unnecessary for females to have the faculty of laying royal eggs; such prodigality of means did not appear to him consistent with nature's laws: therefore he asserts in proper terms that the mother bee does not lay royal eggs in cells prepared to that end: he considers the royal cells only as common ones, enlarged by the bees at the moment when the included form is destined to become a queen; and he adds that, in any case, the royal cell would be too deep for the belly of the mother to reach its bottom to deposit an egg.
I admit that Mr. de Reaumur does not say anywhere that he had seen the queen lay in the royal cell; however he had no doubt of this, and after all my observations I see that he had guessed correctly. It is quite certain that, at certain times of the year, the bees prepare royal cells, that the queens lay eggs in them, and that from these eggs worms hatch which become queens.
Mr. Schirach's objection concerning the length of the royal cells proves nothing:' the queen does not wait till they are finished, to lay in them; she deposits her eggs in them when they are only sketched and have the shape of an acorn cup. This naturalist, dazzled by the brilliancy of his discovery, did not see the whole truth; he was the first to perceive the resource which nature granted to the bees, for repairing the loss of their queen, and too soon persuaded himself that she had provided no other means for the birth of females. His error arose in not observing the bees in sufficiently narrow hives. Had he used hives such as mine he would have found a confirmation of the opinion of Mr. de Reaumur, in every one that he opened in the spring. In that season, which is the season of swarms, hives in good condition are governed by a prolific queen. One finds in such hives royal cells of a different shape from those which the bees build around worker eggs, which they destine to become queens. They are large cells, attached to the comb by a pedicle, and hanging vertically like stalactites; such, in a word, as Mr. de Reaumur has described. The females do not wait, to lay in them, until they are at full size; we have surprised several depositing the egg, when the cell was only as an acorn cup; the workers never lengthen them until the egg has been laid in them; they enlarge them as fast as the worm grows, and close them when it is about to transform into a royal nymph. It is then true that, in spring, the queen deposits, in the royal cells previously prepared, eggs from which insects of her own sort will hatch. Nature has thus provided a twofold means for the multiplication and conservation of the species of bees.
I have the honor, &c.
FITH LETTER
Experiments Proving That There Are, Sometimes, in the Hives, Worker
Bees Which Lay Fertile Egps.
Pregny, August 25, 1791.
Sir, The singular discovery of Mr. Riem, concerning the existence of fertile workers appeared very doubtful to you,10
You have suspected that the eggs, ascribed to workers by this naturalist, were really laid by small queens, which had been mistaken for workers on account of their small size. However you did not state decisively that Mr. Riem was mistaken; and in the letter which you did me the honor of addressing to me, you invited me to seek, through new experiments, whether there are truly in the hives worker bees capable of laying fertile eggs. I have made those experiments, Sir, with great care; you will judge of the degree of confidence which they merit.
On the 5th of August 1788, we found eggs and worms of drones in two of my hives, which had both been deprived of queens for some time. We saw there also the first rudiments of several royal cells, appended like sta-lactites at the edges of the combs. In those cells were some male eggs. As I was perfectly certain that there was no queen of the large size among the bees of these two hives, it was evident that the eggs that were there, the number of which increased every day, had been laid, either by queens of the small size, or by fertile workers. I had reason to believe that it was actually common bees that laid them; for we often observed bees of the latter kind introducing the posterior part of their body into the cells, assuming the same attitude as the queen when she is about to lay. But notwithstanding all our efforts, we had never been able to seize one in this position to examine her more closely; and we did not wish to assert anything until we had been able to hold in our fingers one of the bees that had been laying. We therefore continued our observations with the same assiduity, hoping that, by some fortunate hazard or in a moment of dexterity, we could secure one of those bees. For more than a month, all our endeavors failed.
Burnens then proposed to me to make upon those two hives an operation which required so much courage and patience that I had not dared mention it to him, although the same plan had occurred to me. He proposed to examine, separately, all the bees in these hives, to discover whether some small queen had not insinuated herself among them, which had escaped our first researches. This experiment was very important, for if we did not find any small queens, we would acquire the demonstrative evidence that the eggs whose origin we were seeking had been laid by simple workers.
To accomplish with all possible exactness an operation of this nature, immersing the bees was not sufficient. You know, Sir, that the contact of water stiffens their exterior parts, that it alters to a certain point the shape of their organs; and as the small queens much resemble the workers, the
## Fifth Letter.
**slightest alteration in their shape would not have permitted us to distinguish, with sufficient precision, to which species those that were immersed belonged. Therefore, it was necessary, to take each bee, one at a time, in the hives, alive, in spite of their anger, and observe their specific character with the greatest care. This is what Burnens undertook and executed with inconeivable dexterity. He spent 11 days in this operation, and during the entire time scarcely allowed himself any other relaxation than that required for the relief of his eyes. He took in his hands every one of the bees that composed these two hives, examined carefully their proboscis, their posterior legs, their sting; he found none that did not possess the characters of the common bee; that is to say, the little basket on the posterior legs, the long tongue and the straight sting. Having previously prepared glass cases containing combs, he placed each bee into one of them after having examined her: needless to say that he kept them confined; this precaution was indispensable, for the experiment was not yet ended; it was not sufficient to make sure that they were worker bees, it was also necessary to see whether any of them would lay eggs. So, we inspected, for several days, the combs which we had given them, and we shortly noticed newly laid eggs from which drone worms hatched in due time.**
**Burnens had held between his fingers the bees that produced them, and as he was positive that he had handled only common bees, it was demonstrated that there are, sometimes, in the hives, fertile workers.**
**After having verified Mr. Rlem's discovery by so decisive an experiment, we replaced all the bees that we had examined, into very thin glass hives; these hives which were only an inch-and-a-half thick could contain but a single row of combs; they were thus very favorable for observation. We no longer doubted that, by persisting in watching our bees, we should surprise one of the fertile ones in the act of laying, and seize her. We desired to dissect her, compare the state of her ovaries with those of queens, and take note of the difference. At length, on the 8th of September, we had the good fortune to succeed.**
**A bee was noticed in a cell, in the position of a laying female; we did not give her time to leave the cell; we promptly opened the hive and seized that bee; she had all the external characteristics of common bees; the only difference we could recognize, and a very slight one, was that her belly was smaller and more slender than those of the other workers. We dissected her and found her ovaries smaller, more fragile, composed of a less number of ovludents than the ovaries of queens; the filaments which contained the eggs were exceedingly fine and exhibited slight swellings at equal distances. We counted 11 eggs of sensible size, some of which appeared ready to be laid. This ovary was double, like that of a queen.**
**On the 9th of September, we seized another fertile worker, the instant she laid, and dissected her. Her ovary was still less developed than that of the preceding one, we counted only 4 eggs that had reached maturity. Burnens extracted one of these eggs from the oviduct which held it, and succeeded in fixing it by an end on a strip of glass: we may say in passing that this indicates that it is in the ovluducts themselves that the eggs are coated with the viscous liquid with which they come to light and not in their passage under the spherical sac, as Swammerdam believed.**During the remainder of this month, we found still in the same hives 10 fertile workers, which we also dissected. We distinguished easily the ovaries in most of those bees; in a few, however, we found no trace of them; the oviduets of the latter were apparently but imperfectly developed, and more skill than we had yet acquired in dissecting was necessary for their discovery.
Fertile workers never lay eggs of common bees; they lay only drone eggs. Mr. Riem had already observed this singular fact, and in this regard, all my observations confirm his own. I shall only add to what he says that fertile workers are not absolutely indifferent in the choice of the cells for depositing their eggs. They always prefer to lay them in large ones, and lay in small cells only when they do not find any of a larger diameter; but they have this in common with queens whose foundation has been retarded that they also lay eggs in royal cells.