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diff --git a/26457-8.txt b/26457-8.txt new file mode 100644 index 0000000..8de801e --- /dev/null +++ b/26457-8.txt @@ -0,0 +1,5329 @@ +The Project Gutenberg EBook of New observations on the natural history of +bees, by Francis Huber + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: New observations on the natural history of bees + +Author: Francis Huber + +Translator: Anonymous + +Release Date: August 28, 2008 [EBook #26457] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES *** + + + + +Produced by Louise Pryor, Steven Giacomelli and the Online +Distributed Proofreading Team at https://www.pgdp.net (This +file was produced from images produced by Core Historical +Literature in Agriculture (CHLA), Cornell University) + + + + + +{Transcriber's note + +The spelling in the original is sometimes idiosyncratic. It has not +been changed, but a few obvious errors have been corrected. The +corrections are listed at the end of this etext.} + + +{Illustration: The figures that are referred to in the text} + + + + + NEW OBSERVATIONS + ON THE + NATURAL HISTORY + OF + BEES, + + BY + FRANCIS HUBER. + + TRANSLATED FROM THE ORIGINAL. + + * * * * * + + EDINBURGH: + + PRINTED FOR JOHN ANDERSON, + AND SOLD BY + LONGMAN, HURST, REES, AND ORME, + LONDON. + + ALEX SMELLIE, Printer. + + 1806. + + + + + _To + SIR JOSEPH BANKS, BART._ + + _KNIGHT OF THE MOST HONOURABLE ORDER + OF THE BATH, A PRIVY COUNCILLOR, + PRESIDENT OF THE ROYAL + SOCIETY OF LONDON, + &c. &c._ + + _THIS TRANSLATION + IS INSCRIBED._ + + + + +CONTENTS. + + + LETTER 1.--On the impregnation of the queen bee page 1 + + LETTER 2.--Sequel of observations on the impregnation + of the queen bee 41 + + LETTER 3.--The same subject continued; observations + on retarding the fecundation of queens 44 + + LETTER 4.--On M. Schirach's discovery 76 + + LETTER 5.--Experiments proving that there are + sometimes common bees which lay fertile eggs 89 + + LETTER 6.--On the combats of queens; the massacre + of the males; and what succeeds in a hive + where a stranger queen is substituted for + the natural one 108 + + LETTER 7.--Sequel of observations on the reception + of a stranger queen; M. de Reaumur's + observations on the subject 137 + + LETTER 8.--Is the queen oviparous? What influence + has the size of the cells where the eggs + are deposited on the bees produced? + Researches on the mode of spinning the coccoons 145 + + LETTER 9.--On the formation of swarms 171 + + LETTER 10.--The same subject continued 201 + + LETTER 11.--The same subject continued 223 + + LETTER 12.--Additional observations on queens + that lay only the eggs of drones, and on + those deprived of the antennæ 237 + + LETTER 13.--Economical considerations on bees 253 + + APPENDIX 275 + + + + +TRANSLATOR'S PREFACE + + +The facts contained in this volume are deeply interesting to the +Naturalist. They not only elucidate the history of those industrious +animals, whose nature is the peculiar subject of investigation, but they +present some singular features in physiology which have hitherto been +unknown. + +The industry of bees has proved a fertile source of admiration in all +countries and in every age; and mankind have endeavoured to render it +subservient to their gratifications or emolument. Hence innumerable +theories, experiments, and observations have ensued, and uncommon +patience has been displayed in prosecuting the enquiry. But although +many interesting peculiarities have been discovered, they are so much +interwoven with errors, that no subject has given birth to more +absurdities than investigations into the history of bees: and +unfortunately those treatises which are most easily attained, and the +most popular, only serve to give such absurdities a wider range, and +render it infinitely more difficult to eradicate them. A considerable +portion of the following work is devoted to this purpose. The reader +will judge of the success which results from the experiments that have +been employed. + +Perhaps this is not the proper place to bestow an encomium on a treatise +from which so much entertainment and instruction will be derived. +However, to testify the estimation in which it is held in other nations, +the remarks upon it by the French philosopher Sue, may be quoted, 'The +observations are so consistent, and the consequences seem so just, that +while perusing this work, it appears as if we had assisted the author +in each experiment, and pursued it with equal zeal and interest. Let us +invite the admirers of nature to read these observations; few are equal +to them in excellence, or so faithfully describe the nature, the habits, +and inclinations of the insects of which they treat.' + +It is a remarkable circumstance that the author laboured under a defect +in the organs of vision, which obliged him to employ an assistant in his +experiments. Thus these discoveries may be said to acquire double +authority. But independent of this the experiments are so judiciously +adapted to the purposes in view, and the conclusions so strictly +logical, that there is evidently very little room for error. The talents +of _Francis Burnens_, this philosophic assistant, had long been devoted +to the service of the author, who, after being many successive years in +this manner aided in his researches, was at last deprived of him by some +unfortunate accident. + +Whether the author has prosecuted his investigation farther does not +appear, as no other production of his pen is known in this island. + +It is vain to attempt a translation of any work without being to a +certain degree skilled in the subject of which it treats. Some parts of +the original of the following treatise, it must be acknowledged, are so +confused, and some so minute, that it is extremely difficult to give an +exact interpretation. But the general tenor, though not elegant, is +plain and perspicuous; and such has it been here retained. + + + + +LETTER I. + +_ON THE IMPREGNATION OF THE QUEEN BEE._ + + + SIR, + +When I had the honour at Genthod of giving you an account of my +principal experiments on bees, you desired me to transmit a written +detail, that you might consider them with greater attention. I hasten, +therefore, to extract the following observations from my journal.--As +nothing can be more flattering to me than the interest you take in my +researches, permit me to remind you of your promise to suggest new +experiments{A}. + +After having long studied bees in glass hives constructed on M. de +Reaumur's principle, you have found the form unfavourable to an +observer. The hives being too wide, two parallel combs were made by the +bees, consequently whatever passed between them escaped observation. +From this inconvenience, which I have experienced, you recommended much +thinner hives to naturalists, where the panes should be so near each +other, that only a single row of combs could be erected between them. I +have followed your admonitions, Sir, and provided hives only eighteen +lines in width, in which I have found no difficulty to establish swarms. +However, bees must not be entrusted with the charge of constructing a +single comb: Nature has taught them to make parallel ones, which is a +law they never derogate from, unless when constrained by some particular +arrangement. Therefore, if left to themselves in these thin hives, as +they cannot form two combs parallel to the plane of the hive, they will +form several small ones perpendicular to it, and, in that case, all is +equally lost to the observer. Thus it became essential previously to +arrange the position of the combs. I forced the bees to build them +perpendicular to the horizon, and so that the lateral surfaces were +three or four lines from the panes of the hive. This distance allows the +bees sufficient liberty, but prevents them from collecting in too large +clusters on the surface of the comb. By such precautions, bees are +easily established in very thin hives. There they pursue their labours +with the same assiduity and regularity; and, every cell being exposed, +none of their motions can be concealed. + +It is true, that by compelling these insects to a habitation where they +could construct only a single row of combs, I had, in a certain +measure, changed their natural situation, and this circumstance might +possibly have affected their instinct. Therefore, to obviate every +objection, I invented a kind of hives, which, without losing the +advantages of those very thin, at the same time approached the figure of +common hives where bees form several rows of combs. + +I took several small fir boxes, a foot square and fifteen lines wide, +and joined them together by hinges, so that they could be opened and +shut like the leaves of a book{B}. When using a hive of this +description, we took care to fix a comb in each frame, and then +introduced all the bees necessary for each particular experiment. By +opening the different divisions successively, we daily inspected both +surfaces of every comb. There was not a single cell where we could not +distinctly see what passed at all times, nor a single bee, I may almost +say, with which we were not particularly acquainted. Indeed, this +construction is nothing more than the union of several very flat hives +which may be separated. Bees, in such habitations, must not be visited +before their combs are securely fixed in the frames, otherwise, by +falling out, they may kill or hurt them, as also irritate them to that +degree that the observer cannot escape stinging, which is always +painful, and sometimes dangerous: but they soon become accustomed to +their situation, and in some measure tamed by it; and, in three days, we +may begin to operate on the hive, to open it, remove part of the combs, +and substitute others, without the bees exhibiting too formidable +symptoms of displeasure. You will remember, Sir, that on visiting my +retreat, I shewed you a hive of this kind that had been a long time in +experiment, and how much you were surprised that the bees so quietly +allowed us to open it. + +In these hives, I have repeated all my observations, and obtained +exactly the same results as in the thinnest. Thus, I think, already to +have obviated any objections that may arise concerning the supposed +inconvenience of flat hives. Besides, I cannot regret the repetition of +my labours; by going over the same course several times, I am much more +certain of having avoided error; and it also appears, that some +advantages are found in these which may be called _Book_ or +_Leaf-hives_, as they prove extremely useful in the economical treatment +of bees, which shall afterwards be detailed. + +I now come to the particular object of this letter, the fecundation of +the queen bee; and I shall, in a few words, examine the different +opinions of naturalists on this singular problem. Next I shall state the +most remarkable observations which their conjectures have induced me to +make, and then describe the new experiments by which I think I have +solved the problem{C}. + +Swammerdam, who studied bees with unremitting attention, and who never +could see a real copulation between a drone and a queen, was satisfied +that copulation was unnecessary for fecundation of the eggs: but having +remarked that, at certain times, the drones exhaled a very strong odour, +he thought this odour was an emanation of the _aura seminalis_, or the +_aura seminalis_ itself, which operated fecundation by penetrating the +body of the female. His conjecture was confirmed on dissecting the male +organs of generation; for he was so much struck with the disproportion +between them and those of the female, that he did not believe copulation +possible. His opinion, concerning the influence of the odour, had this +farther advantage, that it afforded a good reason for the prodigious +number of the males. There are frequently fifteen hundred or two +thousand in a hive; and, according to Swammerdam, it is necessary they +should be numerous, that the emanation proceeding from them may have an +intensity or energy sufficient to effect impregnation. + +Though M. de Reaumur has refuted this hypothesis by just and conclusive +reasoning, he has failed to make the sole experiment that could support +or overturn it. This was to confine all the drones of a hive in a tin +case, perforated with minute holes, which might allow the emanation of +the odour to escape, but prevent the organs of generation from passing +through. Then, this case should have been placed in a hive well +inhabited, but completely deprived of males, both of large and small +size, and the consequence attended to. It is evident, had the queen laid +eggs after matters were thus disposed, that Swammerdam's hypothesis +would have acquired probability; and on the contrary it would have been +confuted had she produced no eggs, or only sterile ones. However the +experiment has been made by us, and the queen remained barren; +therefore, it is undoubted, that the emanation of the odour of the males +does not impregnate bees. + +M. de Reaumur was of a different opinion. He thought that the queen's +fecundation followed actual copulation. He confined several drones in a +glass vessel along with a virgin queen: he saw the female make many +advances to the males; but, unable to observe any union so intimate +that it could be denominated copulation, he leaves the question +undecided. We have repeated this experiment: we have frequently confined +virgin queens with drones of all ages: we have done so at every season, +and witnessed all their advances and solicitations to the males: we have +even believed we saw a kind of union between them, but so short and +imperfect that it was unlikely to effect impregnation. Yet, to neglect +nothing, we confined the virgin queen, that had suffered the approaches +of the male, to her hive. During a month that her imprisonment +continued, she did not lay a single egg; therefore, these momentary +junctions do not accomplish fecundation. + +In the _Contemplation de la Nature_, you have cited the observations of +the English naturalist Mr Debraw. They appear correct, and at last to +elucidate the mystery. Favoured by chance, the observer one day +perceived at the bottom of cells containing eggs, a whitish fluid, +apparently spermatic, at least, very different from the substance or +jelly which bees commonly collect around their new hatched worms. +Solicitous to learn its origin, and conjecturing that it might be the +male prolific fluid, he began to watch the motions of every drone in the +hive, on purpose to seize the moment when they would bedew the eggs. He +assures us, that he saw several insinuate the posterior part of the body +into the cells, and there deposit the fluid. After frequent repetition +of the first, he entered on a long series of experiments. He confined a +number of workers in glass bells along with a queen and several males. +They were supplied with pieces of comb containing honey, but no brood. +He saw the queen lay eggs, which were bedewed by the males, and from +which larvæ were hatched, consequently, he could not hesitate advancing +as a fact demonstrated, that male bees fecundate the queen's eggs in +the manner of frogs and fishes, that is, after they are produced. + +There was something very specious in this explanation: the experiments +on which it was founded seemed correct; and it afforded a satisfactory +reason for the prodigious number of males in a hive. At the same time, +the author had neglected to answer one strong objection. Larvæ appear +when there are no drones. From the month of September until April, hives +are generally destitute of males, yet, notwithstanding their absence, +the queen then lays fertile eggs. Thus, the prolific fluid cannot be +required to impregnate them, unless we can suppose that it is necessary +at a certain time of the year, while at every other season it is +useless. + +To discover the truth amidst these facts apparently so contradictory, I +wished to repeat Mr Debraw's experiments, and to observe more precaution +than he himself had done. First, I sought for the fluid, which he +supposes the seminal, in cells containing eggs. Several were actually +found with that appearance; and, during the first days of observation, +neither my assistant nor myself doubted the reality of the discovery. +But we afterwards found it an illusion arising from the reflection of +the light, for nothing like a fluid was visible, except when the solar +rays reached the bottom of the cells. Fragments of the coccoons of +worms, successively hatched, commonly cover the bottom; and, as they are +shining, it may easily be conceived that, when much illuminated, an +illusory effect results from the light. We proved it by the strictest +examination, for no vestiges of a fluid were perceptible when the cells +were detached and cut asunder. + +Though the first observation inspired us with some distrust of Mr +Debraw's discovery, we repeated his other experiments with the utmost +care. On the 6. of August 1787, we immersed a hive, and, with scrupulous +attention, examined the whole bees while in the bath. We ascertained +that there was no male, either large or small; and having examined all +the combs, we found neither male nymph, nor worm. When the bees were +dry, we replaced them all, along with the queen, in their habitation, +and transported them into my cabinet. They were allowed full liberty; +therefore, they flew about, and made their usual collections; but, it +being necessary that no male should enter the hive during the +experiment, a glass tube was adapted to the entrance, of such dimensions +that two bees only could pass at once; and we watched the tube +attentively during the four or five days that the experiment continued. +We should have instantly observed and removed any male that appeared, +that the result of the experiment might be undisturbed, and I can +positively affirm that not one was seen. However, from the first day, +which was the sixth of August, the queen deposited fourteen eggs in the +workers cells; and all these were hatched on the tenth of the same +month. + +This experiment is decisive, since the eggs laid by the queen of a hive +where there were no males, and where it was impossible one could be +introduced, since these eggs, I say, were fertile, it becomes +indubitable that the fluid of the males is not required for their +exclusion. + +Though it did not appear that any reasonable objection could be started +against this conclusion, yet, as I had been accustomed in all my +experiments to seek for the most trifling difficulties that could arise, +I conceived that Mr Debraw's partisans might maintain, that the bees, +deprived of drones, perhaps would search for those in other hives, and +carry the fecundative fluid to their own habitations for depositing it +on the eggs. + +It was easy to appreciate the force of this objection, for all that was +necessary was a repetition of the former experiments, and to confine the +bees so closely to their hives that none could possibly escape. You +very well know, Sir, that these animals can live three or four months +confined in a hive well stored with honey and wax, and if apertures are +left for circulation of the air. This experiment was made on the tenth +of August; and I ascertained, by means of immersion, that no male was +present. The bees were confined four days in the closest manner, and +then I found forty young larvæ. + +I extended the precautions so far as to immerse this hive a second time, +to assure myself that no male had escaped my researches. Each of the +bees was separately examined, and none was there that did not display +its sting. The coincidence of this experiment with the other, proved +that the eggs were not externally fecundated. + +In terminating the confutation of Mr Debraw's opinion, I have only to +explain what led him into error; and that was, his using queens whose +history he was unacquainted with from their origin. When he observed +the eggs produced by a queen, confined along with males, were fertile, +he thence concluded that they had been bedewed by the prolific fluid in +the cells: but to render his conclusion just, he should first have +ascertained that the female never had copulated, and this he neglected. +The truth is, that, without knowing it, he had used, in his experiments, +a queen after she had commerce with the male. Had he taken a virgin +queen the moment she came from the royal cell, and confined her along +with drones in his vessels, the result would have been opposite; for, +even amidst a seraglio of males, this young queen would never have laid, +as I shall afterwards prove. + +The Lusatian observers, and M. Hattorf in particular, thought the queen +was fecundated by herself, without concourse with the males. I shall +here give an abstract of the experiment on which that opinion is +founded.{D} + +M. Hattorf took a queen whose virginity he could not doubt. He excluded +all the males both of the large and small species, and, in several days, +he found both eggs and worms. He asserts that there were no drones in +the hive, during the course of the experiment; but although they were +absent, the queen laid eggs, from which came worms: whence he considers +she is impregnated by herself. + +Reflecting on this experiment, I do not find it sufficiently accurate. +Males pass with great facility from hive to hive; and M. Hattorf took no +precaution that none was introduced into his. He says, indeed, there was +no male, but is silent respecting the means he adopted to prove the +fact. Though he might be satisfied of no large drone being there, still +a small one might have escaped his vigilance, and fecundated the queen. +With a view to clear up the doubt, I resolved to repeat his experiment, +in the manner described, and without greater care or precaution. + +I put a virgin queen into a hive, from which all the males were +excluded, but the bees left at perfect liberty. For several days I +visited the hive, and found new hatched worms in it. Here then is the +same result as M. Hattorf obtained? But before deducing the same +consequence from it, we had to ascertain beyond dispute that no male had +entered the hive. Thus, it was necessary to immerse the bees, and +examine each separately. By this operation, we actually found four small +males. Therefore, to render the experiment decisive, not only was it +requisite to remove all the drones, but also, by some infallible method, +to prevent any from being introduced, which the German naturalist had +neglected. + +I prepared to repair this omission, by putting a virgin queen into a +hive, from which the whole males were carefully removed; and to be +physically certain that none should enter, a glass tube was adapted at +the entrance of such dimensions that the working bees could freely pass +and repass, but too narrow for the smallest male. Matters continued thus +for thirty days, the workers departing and returning performed their +usual labours: but the queen remained sterile. At the expiration of this +time, her belly was equally slender as at the moment of her origin. I +repeated the experiment several times, and always with the same +consequence. + +Therefore, as a queen, rigorously separated from all commerce with the +male, remains sterile, it is evident she cannot impregnate herself, and +M. Hattorf's opinion is ill-founded. + +Hitherto, by endeavouring to confute or verify the conjectures of all +the authors who had preceded me, by new experiments, I acquired the +knowledge of new facts, but these were apparently so contradictory as to +render the solution of the problem still more difficult. While +examining Mr. Debraw's hypothesis, I confined a queen in a hive, from +which all the drones were removed; the queen nevertheless was fertile. +When considering the opinion of M. Hattorf on the contrary, I put a +queen, of whose virginity I was perfectly satisfied, in the same +situation, she remained sterile. + +Embarrassed by so many difficulties, I was on the point of abandoning +the subject of my researches, when at length by more attentive +reflection, I thought these contradictions might arise from experiments +made indifferently on virgin queens, and on those with whose history I +was not acquainted from the origin, and which had perhaps been +impregnated unknown to me. Impressed with this idea, I undertook a new +method of observation not on queens fortuitously taken from the hive, +but on females decidedly in a virgin state, and whose history I knew +from the instant they left the cell. + +From a very great number of hives, I removed all the virgin females, +and substituted for each a queen taken at the moment of her birth. The +hives were then divided into two classes. From the first, I took the +whole males both large and small, and adapted a glass tube at the +entrance, so narrow, that no drone could pass, but large enough for the +free passage of the common bees. In the hives of the second class, I +left all the drones belonging to them, and even introduced more; and to +prevent them from escaping, a glass tube, also too narrow for the males, +was adapted to the entrance of these hives. + +For more than a month, I carefully watched this experiment, made on a +large scale; but much to my surprise, all the queens remained sterile. +Thus it was proved, that queens confined in a hive would continue barren +though amidst a seraglio of males. + +This result induced me to suspect that the females could not be +fecundated in the interior of the hive, and that it was necessary for +them to leave it for receiving the approaches of the male. To ascertain +the fact was easy, by a direct experiment; and as the point is +important, I shall relate in detail what was done by my secretary and +myself on the 29. June 1788. + +Aware, that in summer the males usually leave the hive at the warmest +time of the day, it was natural for me to conclude that if the queens +were also obliged to go out for impregnation, instinct would induce them +to do so at the same time as the males. + +At eleven in the forenoon, we placed ourselves opposite a hive +containing an unimpregnated queen five days old. The sun had shone from +his rising; the air was very warm; and the males began to leave the +hives. We then enlarged the entrance of that which we wished to observe, +and paid great attention to the bees that entered and departed. The +males appeared, and immediately took flight. Soon afterwards, the young +queen appeared at the entrance; at first she did not fly, but brushed +her belly with her hind legs, and traversed the board a little; neither +workers nor males paid any attention to her. At last, she took flight. +When several feet from the hive, she returned, and approached it as if +to examine the place of her departure, perhaps judging this precaution +necessary to recognize it; she then flew away, describing horizontal +circles twelve or fifteen feet above the earth. We contracted the +entrance of the hive that she might not return unobserved, and placed +ourselves in the centre of the circles described in her flight, the more +easily to follow her and observe all her motions. But she did not remain +long in a situation favourable for us, and rapidly rose out of sight. We +resumed our place before the hive; and in seven minutes, the young queen +returned to the entrance of a habitation which she had left for the +first time. Having found no external appearance of fecundation, we +allowed her to enter. In a quarter of an hour she re-appeared; and, +after brushing herself as before, took flight. Then returning to examine +the hive, she rose so high that we soon lost sight of her. Her second +absence was much longer than the first; twenty-seven minutes elapsed +before she came back. We then found her in a state very different from +that in which she was after her first excursion. The sexual organs were +distended by a white substance, thick and hard, very much resembling the +fluid in the vessels of the male, completely similar to it indeed in +colour and consistence{E}. + +But more evidence than mere resemblance was requisite to establish that +the female had returned with the prolific fluid of the males. We allowed +this queen to enter the hive, and confined her there. In two days, we +found her belly swoln; and she had already laid near an hundred eggs in +the worker's cells. + +To confirm our discovery, we made several other experiments, and with +the same success. I shall continue to transcribe my journal. + +On the second of July, the weather being very fine, numbers of males +left the hives. We set at liberty an unimpregnated young queen, eleven +days old, whose hive had always been deprived of males. Having quickly +left the hive, she returned to examine it, and then rose out of sight. +In a few minutes, she returned without any external marks of +impregnation. In a quarter of an hour, she departed again, but her +flight was so rapid that we could scarcely follow her a moment. This +absence continued thirty minutes. On returning, the last ring of the +body was open, and the sexual organs full of the whitish substance +already mentioned. She was then replaced in the hive from which all the +males were excluded. In two days, we found her impregnated. + +These observations at length demonstrate why M. Hattorf obtained results +so different from ours. His queens, though in hives deprived of males, +had been fecundated, and he thence concludes that sexual intercourse is +not requisite for their impregnation. But he did not confine the queens +to their hives, and they had profited by their liberty to unite with the +males. We, on the contrary, have surrounded our queens with a number of +males; but they continued sterile; because the precaution of confining +the males to their hives had also prevented the queens from departing to +seek that fecundation without, which they could not obtain within. + +These experiments were repeated on queens, twenty, twenty-five, and +thirty days old. All became fertile after a single impregnation; +however, we have remarked some essential peculiarities in the fecundity +of those unimpregnated until the twentieth day of their existence; but +we shall defer speaking of the fact until we can present naturalists +with observations sufficiently secure and numerous to merit their +attention: Yet let me add a few words more. Though neither my assistant +nor myself have witnessed the copulation of a queen and a drone, we +think that, after the detail which has just been commenced, no doubt of +it can remain, or of the necessity of copulation to effect impregnation. +The sequel of experiments, made with every possible precaution, appears +demonstrative. The uniform sterility of queens in hives wanting males, +and in those where they were confined along with them; the departure of +these queens from the hives; and the very conspicuous evidence of +impregnation with which they return, are proofs against which no +objections can stand. But we do not despair of being able next spring to +obtain the complement of this proof, by seizing the female at the very +moment of copulation. + +Naturalists have always been very much embarrassed to account for the +number of males found in most hives, and which seem only a burden on the +community, since they fulfil no function. But we now begin to discern +the object of nature in multiplying them to that extent. As fecundation +cannot be accomplished within, and as the queen is obliged to traverse +the expanse of the atmosphere, it is requisite the males should be +numerous that she may have the chance of meeting some one of them. Were +only two or three drones in each hive, there would be little probability +of their departure at the same instant with the queen, or that they +would meet in their excursions; and most of the females would thus +remain sterile. + +But why has nature prohibited copulation within the hives? This is a +secret still unknown to us. It is possible, however, that some +favourable circumstance may enable us to penetrate it in the course of +our observations. Various conjectures may be formed; but at this day we +require facts, and reject gratuitous suppositions. It should be +remembered, that bees do not form the sole republic among insects +presenting a similar phenomenon; female ants are also obliged to leave +the ant-hills previous to fecundation. + +I cannot request, Sir, that you will communicate the reflections which +your genius will excite concerning the facts I have related. This is a +favour to which I am not yet entitled. But as new experiments will +unquestionably occur to you, whether on the impregnation of the queen or +on other points, may I solicit you to suggest them? They shall be +executed with all possible care; and I shall esteem this mark of +friendship and interest as the most flattering encouragement that the +continuance of my labours can receive. + + _Pregny, 13th August 1789._ + + * * * * * + +_Letter from M. Bonnet to M. Huber._ + +You have most agreeably surprised me, Sir, with your interesting +discovery of the impregnation of the queen bee. It was a fortunate idea, +that she left the hive to be fecundated, and your method of ascertaining +the fact was extremely judicious and well adapted to the object in view. + +Let me remind you, that male and female ants copulate in the air; and +that after impregnation the females return to the ant hills to deposit +their eggs. _Contemplation de la Nature, Part II. chap. 22. note 1._ It +would be necessary to seize the instant when the drone unites with the +female. But how remote from the power of the observer are the means of +ascertaining a copulation in the air. If you have satisfactory evidence +that the fluid bedewing the last rings of the female is the same with +that of the male, it is more than mere presumption in favour of +copulation. Perhaps it may be necessary that the male should seize the +female under the belly, which cannot easily be done but in the air. The +large opening at the extremity of the queen, which you have observed in +so particular a condition, seems to correspond to the singular size of +the sexual parts of the male. + +You wish, my dear Sir, that I should suggest some new experiments on +these industrious republicans. In doing so, I shall take the greater +pleasure and interest, as I know to what extent you possess the valuable +art of combining ideas, and of deducing from this combination results +adapted to the discovery of new facts. A few at this moment occur to me. + +It may be proper to attempt the artificial fecundation of a virgin +queen, by introducing a little of the male's prolific fluid with a +pencil, and at the same time observing every precaution to avoid error. +Artificial fecundation, you are aware, has already succeeded in more +than one animal. + +To ascertain that the queen, which has left the hive for impregnation, +is the same that returns to deposit her eggs, you will find it necessary +to paint the thorax with some varnish that resists humidity. It will +also be right to paint the thorax of a considerable number of workers in +order to discover the duration of their life. This is a more secure +method than slight mutilations. + +For hatching the worm, the egg must be fixed almost vertically by one +end near the bottom of the cell. Is it true, that it is unproductive +unless fixed in this manner? I cannot determine the fact; and therefore +leave it to the decision of experiment. + +I formerly mentioned to you that I had long doubted the real nature of +the small ovular substances deposited by queens in the cells, and my +inclination to suppose them minute worms not yet begun to expand. Their +elongated figure seems to favour my suspicions. It would therefore be +proper to watch them with the utmost assiduity, from the instant of +production until the period of exclusion. If the integument bursts, +there can be no doubt that these minute substances are real eggs. + +I return to the mode of operating copulation. The height that the queen +and the males rise to in the air prevent us from seeing what passes +between them. On that account, the hive should be put into an apartment +with a very lofty ceiling. M. de Reaumur's experiment of confining a +queen with several males in a glass vessel, merits repetition; and if, +instead of a vessel, a glass tube, some inches in diameter and several +feet long, were used, perhaps something satisfactory might be +discovered. + +You have had the fortune to observe the small queens mentioned by the +Abbe Needham, but which he never saw. It will be of great importance to +dissect them for the purpose of finding their ovaries. When M. Reims +informed me that he had confined three hundred workers, along with a +comb containing no eggs, and afterwards found hundreds in it, I strongly +recommended that he should dissect the workers. He did so; and informed +me that eggs were found in three. Probably without being aware of it, he +has dissected small queens. As small drones exist, it is not surprising +if small queens are produced also, and undoubtedly by the same external +causes. + +It is of much consequence to be intimately acquainted with this species +of queens, for they may have great influence on different experiments +and embarrass the observer: we should ascertain whether they inhabit +pyramidal cells smaller than the common, or hexagonal ones. + +M. Schirach's famous experiment on the supposed conversion of a common +worm into a royal one, cannot be too often repeated, though the Lusatian +observers have already done it frequently. I could wish to learn +whether, as the discoverer maintains, the experiment will succeed only +with worms, three or four days old, and never with simple eggs. + +The Lusatian observers, and those of the Palatinate, affirm, that when +common bees are confined with combs absolutely void of eggs, they then +lay none but the eggs of drones. Thus, there must be small queens +producing the eggs of males only, for it is evident they must have +produced those supposed to come from workers. But how is it possible to +conceive that their ovaries contain male eggs alone? + +According to M. de Reaumur, the life of chrysalids may be prolonged by +keeping them in a cold situation, such as an ice-house. The same +experiment should be made on the eggs of a queen; on the nymphs of +drones and workers. + +Another interesting experiment would be to take away all the combs +composing the common cells, and leave none but those destined for the +larvæ of males. By this means we should learn whether the eggs of common +worms, laid by the queen in the large cells, will produce large workers. +It is very probable, however, that deprivation of the common cells might +discourage the bees, because they require them for their honey and wax. +Nevertheless, it is likely, by taking away only part of the common +cells, the workers may be forced to lay common eggs in the cells of +drones. + +I should also wish to have the young larvæ gently removed from the royal +cell, and deposited at the bottom of a common one, along with some of +the royal food. + +As the figure of hives has much influence on the respective disposition +of the combs, it would be a satisfactory experiment, greatly to +diversify their shape and internal dimensions. Nothing could be better +adopted to instruct us how bees can regulate their labours, and apply +them to existing circumstances. This may enable us to discover +particular facts which we cannot foresee. + +The royal eggs and those producing drones, have not yet been carefully +compared with the eggs from which workers come. But they ought to be so, +that we may ascertain whether these different eggs have secret +distinctive characteristics. + +The food supplied by the workers to the royal worm, is not the same with +that given to the common worm. Could we not endeavour, with the point of +a pencil, to remove a little of the royal food, and give it to a common +worm deposited in a cell of the largest dimensions? I have seen common +cells hanging almost vertically, where the queen had laid; and these I +should prefer for this experiment. + +Various facts, which require corroboration, were collected in my Memoirs +on Bees; of this number are my own observations. You can select what is +proper, my dear Sir. You have already enriched the history of bees so +much, that every thing may be expected from your understanding and +perseverance. You know the sentiments with which you have inspired the +CONTEMPLATOR OF NATURE. _Genthod, 18. August 1789._ + + +FOOTNOTES: + +{A} All these letters are addressed to the celebrated naturalist M. +Bonnet.--_T._ + +{B} The leaf or book hive consists of twelve vertical frames or boxes, +parallel to each other, and joined together. Fig. 1. the sides, f f. f +g. should be twelve inches long, and the cross spars, f f. g g. nine or +ten; the thickness of these spars an inch, and their breadth fifteen +lines. It is necessary that this last measure should be accurate; a a. a +piece of comb which guides the bees in their work; d. a moveable slider +supporting the lower part; b b. pegs to keep the comb properly in the +frame or box; four are in the opposite side; e e. pegs in the sides +under the moveable slider to support it. + +A book hive, consisting of twelve frames, all numbered, is represented +fig. 2. Between 6 and 7 are two cases with lids, that divide the hive +into two equal parts, and should only be used to separate the bees for +forming an artificial swarm; a a. two frames which shut up the two sides +of the hive, have sliders, b. b. + +The entrance appears at the bottom of each frame. All should be close +but 1 and 12. However it is necessary that they should open at pleasure. + +The hive is partly open, fig. 3. and shews how the component parts may +be united by hinges, and open as the leaves of a book. The two covers +closing up the sides, a. a. + +Fig. 4. is another view of fig. 1. a a. a piece of comb to guide the +bees; b b. pegs disposed so as to retain the comb properly in the frame; +c c. parts of two shelves; the one above is fixed, and keeps the comb in +a vertical position; the under one, which is moveable, supports it +below. + +{C} I cannot insist that my readers, the better to comprehend what is +here said, shall peruse the Memoirs of M. de Reaumur on Bees, and those +of the Lusace Society; but I must request them to examine the extracts +in M. Bonnet's works, tom. 5. 4to edit. and tom. 10. 8vo, where they +will find a short and distinct abstract of all that naturalists have +hitherto discovered on the subject. + +{D} Vide M. Schirach's History of Bees, in a memoir by M. Hattorf, +entitled, _Physical Researches whether the Queen Bee requires +fecundation by Drones?_ + +{E} It will afterwards appear that what we took for the generative +fluid, was the male organs of generation, left by copulation in the body +of the female. This discovery we owe to a circumstance that shall +immediately be related. Perhaps I should avoid prolixity, by suppressing +all my first observations on the impregnation of the queen, and by +passing directly to the experiments that prove she carries away the +genital organs; but in such observations which are both new and +delicate, and where it is so easy to be deceived, I think service is +done to the reader by a candid avowal of my errors. This is an +additional proof to so many others, of the absolute necessity that an +observer should repeat all his experiments a thousand times, to obtain +the certainty of seeing facts as they really exist. + + + + +LETTER II. + +_SEQUEL OF OBSERVATIONS ON THE IMPREGNATION OF THE QUEEN BEE._ + + + SIR, + +All the experiments, related in my preceding letter, were made in 1787 +and 1788. They seem to establish two facts, which had previously been +the subject of vague conjecture: 1. The queen bee is not impregnated of +herself, but is fecundated by copulation with the male. 2. Copulation is +accomplished without the hive, and in the air. + +The latter appeared so extraordinary, that notwithstanding all the +evidence obtained of it, we eagerly desired to take the queen in the +fact; but, as she always rises to a great height, we never could see +what passed. On that account you advised us to cut part off the wings of +virgin queens. We endeavoured to benefit by your advice, in every +possible manner; but to our great regret, when the wings lost much, the +bees could no longer fly; and, by cutting off only an inconsiderable +portion, we did not diminish the rapidity of their flight. Probably +there is a medium, but we were unable to attain it. On your suggestion, +we tried to render their vision less acute, by covering the eyes with an +opaque varnish, which was an experiment equally fruitless. + +We likewise attempted artificial fecundation, and took every possible +precaution to insure success. Yet the result was always unsatisfactory. +Several queens were the victims of our curiosity; and those surviving +remained sterile. Though these different experiments were unsuccessful, +it was proved that queens leave their hives to seek the males, and that +they return with undoubted evidence of fecundation. Satisfied with this, +we could only trust to time or accident for decisive proof of an actual +copulation. We were far from suspecting a most singular discovery, which +we made in July this year, and which affords complete demonstration of +the supposed event, namely, that the sexual organs of the male remain +with the female.{F} + + +FOOTNOTES: + +{F} The remainder of this Chapter chiefly consists of anatomical +details. These may rather be considered an interruption of the +narrative; and the Translator has judged it expedient to transfer them +to an Appendix. + + + + +LETTER III. + +_THE SAME SUBJECT CONTINUED.--OBSERVATIONS ON RETARDING THE FECUNDATION +OF QUEENS._ + + +In my first letter, I remarked, that when queens were prevented from +receiving the approaches of the male until the twenty-fifth or thirtieth +day of their existence, the result presented very interesting +peculiarities. My experiments at that time were not sufficiently +numerous; but they have since been so often repeated, and the result so +uniform, that I no longer hesitate to announce, as a certain discovery, +the singularities which retarded fecundation, produces on the ovaries of +the queen. If she receives the male during the first fifteen days of her +life, she remains capable of laying both the eggs of workers and of +drones; but should fecundation be retarded until the twenty-second day, +her ovaries are vitiated in such a manner that she becomes unfit for +laying the eggs of workers, and will produce only those of drones. + +In June 1787, being occupied in researches relative to the formation of +swarms, I had occasion, for the first time, to observe a queen that laid +none but the eggs of males. When a hive is ready to swarm, I had before +observed, that the moment of swarming is always preceded by a very +lively agitation, which first affects the queen, is then communicated to +the workers, and excites such a tumult among them, that they abandon +their labours, and rush in disorder to the outlets of the hive. I then +knew very well the cause of the queen's agitation, and it is described +in the history of swarms, but I was ignorant how the delirium +communicated to the workers; and this difficulty interrupted my +researches. I therefore thought of investigating, by direct experiments, +whether at all times, when the queen was greatly agitated, even not in +the time of the hive swarming, her agitation would in like manner be +communicated to the workers. The moment a queen was hatched, I confined +her to the hive by contracting the entrances. When assailed by the +imperious desire of union with the males, I could not doubt that she +would make great exertions to escape, and that the impossibility of it +would produce a kind of delirium. I had the patience to observe this +queen thirty-four days. Every morning about eleven o'clock, when the +weather was fine and the sunshine invited the males to leave their +hives, I saw her impetuously traverse every corner of her habitation, +seeking to escape. Her fruitless efforts threw her into an uncommon +agitation, the symptoms of which I shall elsewhere describe, and all the +common bees were affected by it. As she never was out all this time, she +could not be impregnated. At length, on the thirty-sixth day, I set her +at liberty. She soon took advantage of it; and was not long of returning +with the most evident marks of fecundation. + +Satisfied with the particular object of this experiment, I was far from +any hopes that it would lead to the knowledge of another very remarkable +fact; how great was my astonishment, therefore, on finding that this +female, which, as usual, began to lay forty-six hours after copulation, +laid the eggs of drones, but none of workers, and that she continued +ever afterwards to lay those of drones only. + +At first, I exhausted myself with conjectures on this singular fact; the +more I reflected on it, the more did it seem inexplicable. At length, by +attentively meditating on the circumstances of the experiment it +appeared there were two principles, the influence of which I should +first of all endeavour to appreciate separately. On the one hand, this +queen had suffered long confinement; on the other, her fecundation had +been extremely retarded. You know, Sir, that queens generally receive +the males about the fifth or sixth day, and this queen had not copulated +until the thirty-sixth. Little weight could be given to the supposition, +that the peculiarity could be occasioned by confinement. Queens, in the +natural state, leave their hives only once to seek the males. All the +rest of their life they remain voluntary prisoners. Thus, it was +improbable that captivity could produce the effect I wished to explain. +At the same time, as it was essential to neglect nothing in a subject so +new, I wished to ascertain whether it was owing to the length of +confinement, or to retarded fecundation. + +Investigating this was no easy matter. To discover whether captivity, +and not retarded fecundation, vitiated the ovaries, it was necessary to +allow a female to receive the approaches of a male, and also to keep her +imprisoned. Now this could not be, for bees never copulate in hives. On +the same account, it was impossible to retard the copulation of a queen +without keeping her in confinement. I was long embarrassed by the +difficulty. At length, I contrived an apparatus, which, though +imperfect, nearly fulfilled my purpose. + +I put a queen, at the moment of her last metamorphosis, into a hive well +stored, and sufficiently provided with workers and males; the entrance +was contracted so as to prevent her exit, but allowed free passage to +the workers. I also made another opening for the queen, and adapted a +glass tube to it, communicating with a cubical glass box eight feet +high. Hither the queen could at all times come and fly about, enjoying a +purer air than was to be found within the hive; but she could not be +fecundated; for though the males flew about within the same bounds, the +space was too limited to admit of any union between them. By the +experiments related in my first letter, copulation takes place high in +the air only: therefore, in this apparatus, I found the advantage of +retarding fecundation, while the liberty the queen now had, did not +render her situation too remote from the natural state. I attended to +the experiment fifteen days. Every fine morning, the young captive left +her hive; she traversed her glass prison, and flew much about, and with +great facility. She laid none during this interval, for she had not +united with a male. On the sixteenth day, I set her at liberty: she left +the hive, rose aloft in the air, and soon returned with full evidence of +impregnation. In two days, she laid, first the eggs of workers, and +afterwards as many as the most fertile queens. + +It thence followed, 1. That captivity did not alter the organs of +queens. 2. When fecundation took place within the first sixteen days, +she produced both species of eggs. + +This was an important experiment. It rendered my labours much more +simple, by clearly pointing out the method to be pursued: it absolutely +precluded the supposed influence of captivity; and left nothing for +investigation but the consequences of retarded fecundation. + +With this view, I repeated the experiment; but, instead of giving the +virgin queen liberty on the sixteenth day, I retained her until the +twenty-first. She departed, rose high in the air, was fecundated, and +returned. Thirty-six hours afterwards, she began to lay: but it was the +eggs of males only, and, although very fruitful afterwards, she laid no +other kind. + +I occupied myself the remainder of 1787, and the two subsequent years, +with experiments on retarded fecundation, and had constantly the same +results. It is undoubted, therefore, that when the copulation of queens +is retarded beyond the twentieth day, only an imperfect impregnation is +operated: instead of laying the eggs of workers and males equally, they +will lay none but those of males. + +I do not aspire to the honour of explaining this singular fact. When the +course of my experiments led me to observe that some queens laid only +the eggs of drones, it was natural to investigate the proximate cause of +such a singularity; and I ascertained that it arose from retarded +fecundation. My evidence is demonstrative, for I can always prevent +queens from laying the eggs of workers, by retarding their fecundation +until the twenty-second or twenty-third day. But, what is the remote +cause of this peculiarity; or, in other words, why does the delay of +impregnation render queens incapable of laying the eggs of workers? This +is a problem on which analogy throws no light: nor in all physiology am +I acquainted with any fact that bears the smallest similarity. + +The problem becomes still more difficult by reflecting on the natural +state of things, that is when fecundation has not been delayed. The +queen then lays the eggs of workers forty-six hours after copulation, +and continues for the subsequent eleven months to lay these alone: and +it is only after this period that a considerable and uninterrupted +laying of the eggs of drones commences. When, on the contrary, +impregnation is retarded after the twentieth day, the queen begins, from +the forty-sixth hour, to lay the eggs of males, and no other kind during +her whole life. As, in the natural state, she lays the eggs of workers +only, during the first eleven months, it is clear that these, and the +male eggs, are not indiscriminately mixed in the oviducts. Undoubtedly +they occupy a situation corresponding to the principles that regulate +laying: the eggs of workers are first, and those of drones behind them. +Farther, it appears that the queen can lay no male eggs until those of +workers, occupying the first place in the oviducts, are discharged. Why, +then, is this order inverted by retarded copulation? How does it happen +that all the workers eggs which the queen ought to lay, if fecundation +was in due time, now wither and disappear, yet do not, impede the +passage of the eggs of drones, which occupy only the second place in the +ovaries. Nor is this all. I have satisfied myself that a single +copulation is sufficient to impregnate the whole eggs that a queen will +lay in the course of at least two years. I have even reason to think, +that a single copulation will impregnate all the eggs that she will lay +during her whole life: but I want absolute proof for more than two +years. This, which is truly a very singular fact in itself, renders the +influence of retarded fecundation still more difficult to be accounted +for. Since a single copulation suffices, it is clear that the male fluid +acts from the first moment on all the eggs that the queen will lay in +two years. It gives them, according to your principles, that degree of +_animation_ that afterwards effects their successive expansion. Having +received the first impressions of life, they grow, they mature, so to +speak, until the day they are laid: and as the laws of laying are +constant, because the eggs of the first eleven months are always those +of workers, it is evident that those which appear first are also the +eggs that come soonest to maturity. Thus, in the natural state, the +space of eleven months is necessary for the male eggs to acquire that +degree of increment they must have attained when laid. This consequence, +which to me seems immediate, renders the problem insoluble. How can the +eggs, which should grow slowly for eleven months, suddenly acquire their +full expansion in forty-eight hours, when fecundation has been retarded +twenty-one days, and by the effect of this retardation alone? Observe, I +beseech you, that the hypothesis of successive expansion is not +gratuitous; it rests on the principles of sound philosophy. Besides, for +conviction that it is well founded, we have only to look at the figures +given by Swammerdam of the ovaries of the queen bee. There we see eggs +in that part of the oviducts contiguous to the vulva, much farther +advanced, and larger than those contained in the opposite part. +Therefore the difficulty remains in full force: it is an abyss where I +am lost. + +The only known fact bearing any relation to that now described, is the +state of certain vegetable seeds, which, although extremely well +preserved, lose the faculty of germination from age. The eggs of workers +may also preserve, only for a very short time, the property of being +fecundated by the seminal fluid; and, after this period, which is about +fifteen or eighteen days, become disorganised to that degree, that they +can no longer be animated by it. I am sensible that the comparison is +very imperfect; besides, it explains nothing, nor does it even put us +on the way of making any new experiments. I shall add but one reflection +more. + +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. + +I had a glass hive constructed of two stages; the higher was filled with +combs of large cells, and the lower with those of common ones. A kind of +division, or diaphraghm, separated these two stages from +each other, having at each side an opening for the passage of the +workers from one stage to the other, but too narrow for the queen. I +put a considerable number of bees into this hive; and, in the upper +part, confined a very fertile queen that had just finished her great +laying of male eggs; therefore she had only those of workers to lay, and +she was obliged to deposit them in the surrounding large cells from the +want of others. My object in this arrangement will already be +anticipated. My reasoning was simple. If the queen laid workers' eggs in +the large cells, and the bees were charged with transporting them if +misplaced, they would infallibly take advantage of the liberty allowed +to pass from either stage: they would seek the eggs deposited in the +large cells, and carry them down to the lower stage containing the cells +adapted for that species. If, on the contrary, they left the common eggs +in the large cells, I should obtain certain proof that they had not the +charge of transporting them. + +The result of this experiment excited my curiosity extremely. We +observed the queen several days without intermission. During the first +twenty-four hours, she persisted in not laying a single egg in the +surrounding cells; she examined them one after another, but passed on +without insinuating her belly into one. She was restless, and traversed +the combs in all directions: her eggs appeared an oppressive burden, but +she persisted in retaining them rather than they should be deposited in +cells of unsuitable diameter. The bees, however, did not cease to pay +her homage, and treat her as a mother. I was amused to observe, when she +approached the edges of the division separating the two stages, that she +gnawed at them to enlarge the passage: the workers approached her, and +also laboured with their teeth, and made every exertion 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, and fell +at random. Then we conceived that the bees would convey them into the +small cells of the lower stage, and we sought them there with the utmost +assiduity; but I can safely affirm there was not one. The eggs that the +queen still laid the third day disappeared as the first. We again sought +them in the small cells, but none were there. The fact is, they are ate +by the workers; and this is what has deceived the naturalists, who +supposed them carried away. They have observed the misplaced eggs +disappear, and, without farther investigation, have asserted that the +bees convey them elsewhere: they take them, indeed, not to convey them +any where, but to devour them. Thus nature has not charged bees with the +care of placing the eggs in the cells appropriated for them, but she has +inspired females themselves with sufficient instinct to know the species +of eggs they are about to lay, and to deposit them in suitable cells. +This has already been observed by M. de Reaumur, and here my +observations correspond with his. Thus it is certain that in the natural +state, when fecundation takes place at the proper time, and the queen +has suffered from nothing, she is never deceived in the choice of the +cells where her eggs are to be deposited; she never fails to lay those +of workers in small cells, and those of males in large ones. The +distinction is important, for the same certainty of instinct is no +longer conspicuous in the conduct of those females whose impregnation +has been deferred. I was oftener than once deceived respecting the eggs +that such queens laid, for they were deposited indiscriminately in small +cells and those of drones; and not aware of their instinct having +suffered, I conceived that the eggs in small cells would produce +workers; therefore I was very much surprised, when, at the moment they +should have been hatched, the bees closed up the cells, and +demonstrated, by anticipation, that the included worms would change into +drones; they actually became males; those produced in small cells were +small, those in large cells large. Thus I must warn observers, who would +repeat my experiments on queens that lay only the eggs of males, not to +be deceived by these circumstances, and expect that eggs of males will +be deposited in the workers cells. + +It is a singular fact, that the females, whose fecundation has been +retarded, sometimes lay the eggs of males in royal cells. I shall prove, +in the history of swarms, that immediately when queens, in the natural +state, begin their great laying of male eggs, the workers construct +numerous royal cells. Undoubtedly, there is some secret relation between +the appearance of male eggs and the construction of these cells; for it +is a law of nature from which bees never derogate. It is not surprising, +therefore, that such cells are constructed in hives governed by queens +laying the eggs of males only. It is no longer extraordinary that these +queens deposit in the royal cells, eggs of the only species they can +lay, for in general their instinct seems affected. But what I cannot +comprehend is, why the bees take exactly the same care of the male eggs +deposited in royal cells, as of those that should become queens. They +provide them more plentifully with food, they build up the cells as if +containing a royal worm; in a word, they labour with such regularity +that we have frequently been deceived. More than once, in the firm +persuasion of finding royal nymphs, we have opened the cells after they +were sealed, yet the nymph of a drone always appeared. Here the instinct +of the workers seemed defective. In the natural state, they can +accurately distinguish the male worms from those of common bees, as they +never fail giving a particular covering to the cells containing the +former. Why then can they no longer distinguish the worms of drones when +deposited in the royal cells? The fact deserves much attention. I am +convinced that to investigate the instinct of animals, we must +carefully observe where it appears to err. + +Perhaps I should have begun this letter with an abstract of the +observations of prior naturalists, on queens laying none but the eggs of +males; however, I shall here repair the omission. + +In a work, _Histoire de la Reine des Abeilles_, translated from the +German by _Blassiere_, there is printed a letter from M. Schirach to +you, dated 15 April 1771, where he speaks of some hives, in which the +whole brood changed into drones. You will remember that he ascribes this +circumstance to some unknown vice in the ovaries of the queen; but he +was far from suspecting that retarded fecundation had been the cause of +vitiation. He justly felicitated himself on discovering a method to +prevent the destruction of hives in this situation, which was simple, +for it consisted in removing the queen that laid the eggs of males only, +and substituting one for her whose ovaries were not impaired. But to +make the substitution effectual, it was necessary to procure queens at +pleasure; a secret reserved for M. Schirach, and of which I shall speak +in the following letter. You observe that the whole experiments of the +German naturalist tended to the preservation of the hives whose queens +laid none except male eggs; and that he did not attempt to discover the +cause of the vice evident in their ovaries. + +M. de Reaumur also says a few words, somewhere, of a hive containing +many more drones than workers, but advances no conjectures on the cause. +However, he adds, as a remarkable circumstance, that the males were +tolerated in this hive until the subsequent spring. It is true that bees +governed by a queen laying only male eggs, or by a virgin queen, +preserve their drones several months after they have been massacred in +other hives. I can ascribe no reason for it, but it is a fact I have +several times witnessed during my long course of observations on +retarded impregnation. In general it has appeared that while the queen +lays male eggs, bees do not massacre the males already perfect in the +hive. PREGNY, _21. August 1791_. + + +FOOTNOTES: + +{G} The experiments suggested in this paragraph, recall a singular +reflection of M. de Reaumur. Where treating of oviparous flies, he says, +it would not be impossible for a hen to produce a living chicken, if, +after fecundation, the eggs she should first lay could by any means be +retained twenty-one days in the oviducts. _Mem. sur. les Insect. tom. 4. +mem. 10._ + + + + +LETTER IV. + +_ON M. SCHIRACH'S DISCOVERY._ + + +When you found it necessary, Sir, in the new edition of your works, to +give an account of M. Schirach's beautiful experiments on the conversion +of common worms into royal ones, you invited naturalists to repeat them. +Indeed such an important discovery required the confirmation of several +testimonies. For this reason, I hasten to inform you that all my +researches establish the reality of the discovery. During ten years +that I have studied bees, I have repeated M. Schirach's experiment so +often, and with such uniform success, that I can no longer have the +least doubt on the subject. Therefore, I consider it an established +fact, when bees lose their queen, and several workers' worms are +preserved in the hive, they enlarge some of their cells, and supply them +not only with a different kind of food, but a greater quantity of it, +and the worms reared in this manner, instead of changing to common bees, +become real queens. I request my readers to reflect on the explanation +you have given of so uncommon a fact, and the philosophical consequences +you have deduced from it. _Contemplation de la Nature, part. II, chap. +27._ + +In this letter I shall content myself with some account of the figure of +the royal cells constructed by bees around those worms that are destined +for the royal state, and terminate with discussing some points wherein +my observations differ from those of M. Schirach. + +Bees soon become sensible of having lost their queen, and in a few hours +commence the labour necessary to repair their loss. First, they select +the young common worms, which the requisite treatment is to convert into +queens, and immediately begin with enlarging the cells where they are +deposited. Their mode of proceeding is curious; and the better to +illustrate it, I shall describe the labour bestowed on a single cell, +which will apply to all the rest, containing worms destined for queens. +Having chosen a worm, they sacrifice three of the contiguous cells: +next, they supply it with food, and raise a cylindrical inclosure +around, by which the cell becomes a perfect tube, with a rhomboidal +bottom; for the parts forming the bottom are left untouched. If the bees +damaged it, they would lay open three corresponding cells on the +opposite surface of the comb, and, consequently, destroy their worms, +which would be an unnecessary sacrifice, and Nature has opposed it. +Therefore, leaving the bottom rhomboidal, they are satisfied with +raising a cylindrical tube around the worm, which, like the other cells +in the comb, is horizontal. But this habitation remains suitable to the +worm called to the royal state only during the first three days of its +existence: another situation is requisite for the other two days it is a +worm. Then, which is so small a portion of its life, it must inhabit a +cell nearly of a pyramidal figure, and hanging perpendicularly; we may +say the workers know it; for, after the worm has completed the third +day, they prepare the place to be occupied by its new lodging. They gnaw +away the cells surrounding the cylindrical tube, mercilessly sacrifice +their worms, and use the wax in constructing a new pyramidal tube, which +they solder at right angles to the first, and work it downwards. The +diameter of this pyramid decreases insensibly from the base, which is +very wide, to the point. During the two days that it is inhabited by +the worm, a bee constantly keeps its head more or less inserted into the +cell, and, when this worker quits it, another comes to occupy its place. +In proportion as the worm grows, the bees labour in extending the cell, +and bring food, which they place before its mouth, and around its body, +forming a kind of cord around it. The worm, which can move only in a +spiral direction, turns incessantly to take the food before its head: it +insensibly descends, and at length arrives at the orifice of the cell. +Now is the time of transformation to a nymph. As any farther care is +unnecessary, the bees close the cell with a peculiar substance +appropriated for it, and there the worm undergoes both its +metamorphoses. + +Though M. Schirach supposes that none but worms three days old are +selected for the royal treatment, I am certain of the contrary; and that +the operation succeeds equally well on those of two days only. I must +be permitted to relate at length the evidence I have of the fact, which +will both demonstrate the reality of common worms being converted into +queens, and the little influence which their age has on the effect of +the operation. + +I put some pieces of comb, with some workers eggs, in the cells, and of +the same kind as those already hatched, into a hive deprived of the +queen. The same day several cells were enlarged by the bees, and +converted into royal cells, and the worms supplied with a thick bed of +jelly. Five were then removed from those cells, and five common worms, +which, forty-eight hours before we had seen come from the egg +substituted for them. The bees did not seem aware of the change; they +watched over the new worms the same as over those chosen by themselves; +they continued enlarging the cells, and closed them at the usual time. +When they had hatched on them seven days{H}, we removed the cells to +see the queens that were to be produced. Two were excluded, almost at +the same moment, of the largest size, and well formed in every respect. +The term of the other cells having elapsed, and no queen appearing, we +opened them. In one, was a dead queen, but still a nymph; the other two +were empty. The worms had spun their silk coccoons, but died before +passing into their nymphine state, and presented only a dry skin. I can +conceive nothing more conclusive than this experiment. It demonstrates +that bees have the power of converting the worms of workers into queens; +since they succeeded in procuring queens, by operating on the worms +which we ourselves had selected. It is equally demonstrated, that the +success of the operation does not depend on the worms being three days +old, as those entrusted to the bees were only two. Nor is this all; bees +can convert worms still younger into queens. The following experiment +showed, that when the queen is lost, they destine worms only a few +hours old to replace her. + +I was in possession of a hive, which being long deprived of the female, +had neither egg nor worm. I provided a queen of the greatest fertility; +and she immediately began laying in the cells of workers. I removed this +female before being quite three days in the hive, and before any of her +eggs were hatched. The following morning, that is, the fourth day, we +counted fifty minute worms, the oldest scarcely hatched twenty-four +hours. However, several were already destined for queens, which was +proved by the bees depositing around them a much more abundant provision +of food than is supplied to common worms. Next day, the worms were near +forty hours old: the bees had enlarged and converted their hexagonal +cells into cylindrical ones of the greatest capacity. During the +subsequent days, they still laboured at them, and closed them on the +fifth from the origin of the worms. Seven days after sealing of the +first of these royal cells, a queen of the largest size proceeded from +it. She immediately rushed towards the other royal cells, and +endeavoured to destroy their nymphs and worms. In another letter, I +shall recount the effects of her fury. + +From these details, you will observe, Sir, that M. Schirach's +experiments had not been sufficiently diversified when he affirmed that +it was essential for the conversion of common worms into queens, they +should be three days old. It is undoubted, that equal success attends +the experiment not only with worms two days old, but also when they have +been only a few hours in existence. + +After my researches to corroborate M. Schirach's discovery, I was +desirous of learning whether, as this observer conceives, the only means +which the bees have of procuring a queen, is giving the common worms a +certain kind of aliment, and rearing them in the largest cells. You +will remember, that M. de Reaumur's sentiments are very different: "The +mother should lay, and she does lay, eggs from which flies fit for being +mothers must in their turn proceed. She does so; and it is evident the +workers know what she is to do. Bees, to which the mother is so +precious, seem to take a peculiar interest in the eggs that one is to +proceed from, and to consider them of the greatest value. They construct +particular cells where they are to be deposited.--The figure of a royal +cell only begun, very much resembles a cup, or, more correctly speaking, +the cup that has lost its acorn." + +M. de Reaumur, though he did not suspect the possibility of a common +worm being converted into a queen, conceived that the queen bee laid a +particular species of eggs in the royal cells, from which worms should +come that would be queens. According to M. Schirach, on the other hand, +bees always having the power of procuring a queen by bringing up worms +three days old in a particular manner, it would be needless for nature +to grant females the faculty of laying royal eggs. Such prodigality is, +in his eyes, inconsistent with the ordinary laws of nature. Therefore he +maintains, in direct terms, that she does not lay royal eggs in cells +purposely prepared to receive them. He considers the royal cells only as +common ones, enlarged by the bees at the moment when the included worm +is destined for a queen; and adds, that the royal cell would always be +too long for the belly of the mother to reach the bottom. + +I admit that M. de Reaumur no where says he has seen the queen lay in +the royal cell. However he did not doubt the fact; and, after all my +observations, I must esteem his opinion just. It is quite certain that, +at particular periods of the year, the bees prepare royal cells; that +the females deposit their eggs in them; and that worms, which shall +became queens, proceed from these eggs. + +M. Schirach's objection, concerning the length of the cells, proves +nothing; for the queen does not delay depositing her egg till they are +finished. While only sketched and shaped like the cup of an acorn, she +lays it. This naturalist, dazzled by the brilliancy of his discovery, +saw only part of the truth. He was the first to find out the resource +granted to bees by nature, for repairing the loss of their queen; and +too soon persuaded himself that she had provided no other resource for +the production of females. This error arose from not observing bees in +very flat hives: had he used such as mine, he would have found, on +opening them in spring, a confirmation of M. de Reaumur's opinion. Then, +which is the season of swarming, hives in good condition are governed by +a very fruitful queen: there are royal cells of a figure widely +different from those constructed around the worms destined by the bees +for queens. They are large, attached to the comb by a stalk, and +hanging vertically like stalactites, such, in short, as M. de Reaumur +has described them. The females lay in them before completion. We have +surprised a queen depositing the egg when the cell was only as the cup +of an acorn. The workers never lengthen them until the egg has been +laid. In proportion as the worm grows, they are enlarged, and closed by +the bees when the first transformation approaches. Thus it is true, +that, in spring, the queen deposits in royal cells, previously prepared, +eggs from which flies of her own species are to come. Nature has, +therefore, provided a double means for the multiplication and +conservation of their race. + + _PREGNY, 24. August 1791._ + + +FOOTNOTES: + +{H} The author's meaning here is obscure.--T. + + + + +LETTER V. + +_EXPERIMENTS PROVING THAT THERE ARE SOMETIMES COMMON BEES WHICH LAY +FERTILE EGGS._ + + +The singular discovery of M. Riems, concerning the existence of fertile +workers, has appeared very doubtful to you, Sir. You have suspected that +the eggs ascribed to workers by this naturalist had actually been +produced by small queens, which, on account of their size, were +confounded with common bees. But you do not positively insist that M. +Riems is deceived; and, in the letter which you did me the honour to +address to me, you requested me to investigate, by new experiments, +whether there are actually working bees capable of laying fertile eggs. +I have made these experiments with great care: and it is for you to +judge of the confidence they merit. + +On the fifth of August 1788, we found the eggs and worms of large drones +in two hives, which had both been some time deprived of queens. We also +observed the rudiments of some royal cells appended like stalactites to +the edges of the combs. The eggs of males were in them. Being perfectly +secure that there was no queen of large size among the bees of these two +hives, the eggs, which daily became more numerous, were evidently laid +either by queens of small size or by fertile workers. I had reason to +believe it was actually by common bees, for we had frequently observed +them inserting the posterior part into the cells; and assuming the same +attitude as the queen when laying. But, not withstanding every exertion, +we had never been able to seize one in this situation, to examine it +more narrowly. And we were unwilling to assert any thing positively, +without having the bees in our hands that had actually laid. Therefore +our observations were continued with equal assiduity, in hopes that, by +some fortunate chance, or in a moment of address, we could secure one of +them. More than a month all our endeavours were abortive. + +My assistant then offered to perform an operation that required both +courage and patience, and which I could not resolve to suggest, though +the same expedient had occurred to myself. He proposed to examine each +bee in the hive separately, to discover whether some small queen had not +insinuated herself among them, and escaped our first researches. This +was an important experiment; for, should no small queen be found, it +would be demonstrative evidence that the eggs had been laid by simple +workers. + +To perform this operation with all possible exactness, immersing the +bees was not enough. You know, Sir, that the contact of water stiffens +their organs, that it produces a certain alteration of their external +figure: and, from the resemblance of small queens to workers, the +slightest alteration of shape would prevent us from distinguishing with +sufficient accuracy to what species those immersed might belong. +Therefore it was necessary to seize the whole bees of both hives, +notwithstanding their irritation, and examine their specific character +with the utmost care. This my assistant undertook, and executed with +great address. Eleven days were employed in it; and, during all that +time, he scarcely allowed himself any relaxation, but what the relief of +his eyes required. He took every bee in his hand; he attentively +examined the trunk, the hind limbs, and the sting: there was not one +without the characteristics of the common bee, that is, the little +basket on the hind legs, the long trunk, and the straight sting. He had +previously prepared glass cases containing combs. Into these, he put +each bee after examination. It is superfluous to observe they were +confined, which was a precaution indispensible until termination of the +experiment. Neither was it enough to establish that the whole were +workers; we had also to continue the experiment, and observe whether any +would produce eggs. Thus we examined the cells for several days, and +soon observed new laid eggs, from which the worms of drones came at the +proper time. My assistant held in his hands the bees that produced them; +and as he was perfectly certain they were common ones, it is proved that +there are sometimes fertile workers in hives. + +Having ascertained M. Schirach's discovery, by so decisive an +experiment, we replaced all the bees examined, in very thin glass hives, +being only eighteen lines thick, and capable of containing but a single +row of combs, and thus were extremely favourable to the observer. We +thought, by strictly persisting to watch the bees, we might surprise a +fertile one in the act of laying, seize and dissect her. This we were +desirous of doing, for the purpose of comparing her ovaries with those +of queens, and to ascertain the difference. At length, on the eighth of +September, we had the good fortune to succeed. + +A bee appeared in the position of a female laying. Before she had time +to leave the cell, we suddenly opened the hive and seized her. She +presented all the external characteristics of common bees; the only +difference we could recognise, and that was a very slight one, consisted +in the belly seeming less and more slender than that of workers. On +dissection, her ovaries were found more fragile, smaller and composed of +fewer oviducts than the ovaries of queens. The filaments containing the +eggs were extremely fine, and exhibited swellings at equal distances. We +counted eleven eggs of sensible size, some of which appeared ripe for +laying. This ovary was double like that of queens. + +On the ninth of September, we seized another fertile worker the instant +she laid, and dissected her. The ovary was still less expanded than that +of the preceding bee, and only four eggs had attained maturity. My +assistant extracted one from the oviducts, and succeeded in fixing it by +an end on a glass slider. We may take this opportunity of remarking, +that it is in the oviducts themselves the eggs are imbued with the +viscous liquid, with which they are produced, and not in passing through +the spherical sac as Swammerdam believed. During the remainder of this +month, we found ten fertile workers in the same hives, and dissected +them all. In most, the ovaries were easily distinguished, but in some we +could not discern the faintest traces of them. In these last, the +oviducts to all appearance were but imperfectly developed, and more +address than we had acquired in dissection was necessary to distinguish +them. + +Fertile workers never lay the eggs of common bees; they produce none but +those of males. M. Riems had already observed this singular fact; and +here all my observations correspond with his. I shall only add to what +he says, that fertile workers are not absolutely indifferent in the +choice of cells for depositing their eggs. They always prefer large +ones; and only use small cells when unable to find those of larger +diameter. But they so far correspond with queens whose impregnation has +been retarded, that they sometimes lay in royal cells. + +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. + +I could no longer doubt the truth of my conjecture; at the same time, I +know not whether the truth will appear as rigorous to you, Sir, as it +does to myself. But I reason in the following manner: If it is certain +that fertile workers are always produced in the vicinity of royal cells, +it is no less true, that in itself, the vicinity is indifferent; for the +size and figure of these cells can produce no effect on the worms in +those surrounding them; there must be something more; we know that a +particular aliment is conveyed to the royal cells; we also know, that +this aliment has a very powerful effect on the ovaries; that it alone +can unfold the germ. Thus, we must necessarily suppose the worms in the +adjacent cells have had a portion of the same food. This is what they +gain, therefore, by vicinity to the royal cells. The bees, in their +course thither, will pass in numbers over them, stop and drop some +portion of the jelly destined for the royal larvæ. This reasoning, I +presume, is consistent with the principles of sound logic. + +I have repeated the experiment now described so often, and weighed all +the concomitant circumstances with so much care, that whenever I please, +I can produce fertile workers in my hives. The method is simple. I +remove the queen from a hive; and very soon the bees labour to replace +her, by enlarging several cells, containing the brood of workers, and +supplying the included worms with the royal jelly. Portions of this +aliment also fall on the young larvæ deposited in the adjacent cells, +and it unfolds the ovaries to a certain degree. Fertile workers are +constantly produced in hives where the bees labour to replace their +queen; but we very rarely find them, because they are attacked and +destroyed by the young queens reared in the royal cells. Therefore, to +save them, all their enemies must be removed, and the larvæ of the royal +cells taken away before undergoing their last metamorphoses. Then the +fertile workers, being without rivals at the time of their origin, will +be well received, and, by taking the precaution to mark them, it will be +seen, in a few days, that they produce the eggs of males. Thus, the +whole secret of this proceeding consists in removing the royal cells at +the proper time; that is, after being sealed, and previous to the young +queens leaving them{K}. + +I shall add but a few words to this long letter. There is nothing so +very surprising in the production of fertile workers, when we consider +the consequences of M. Shirach's beautiful discovery. But why do they +lay male eggs only? I can conceive, indeed, that the reason of their +laying few is from their ovaries being but imperfectly expanded, but I +can form no idea why all the eggs should be those of males, neither can +I any better account for their use in hives; and hitherto, I have made +no experiments on their mode of fecundation. + + _PREGNY, 25. August 1791._ + + +FOOTNOTES: + +{I} It is difficult to discover whether the author thinks, as some +naturalists, that bees are instrumental in hatching the eggs.--T. + +{J} The original is extremely confused in the preceding passages.--T. + +{K} I have frequently seen queens, at the moment of production, begin +first by attacking the royal cells and then the common ones beside them. +As I had not seen fertile workers when I first observed this fact, I +could not conceive from what motive the fury of the queen was thus +directed towards the common cells. But now I know they can distinguish +the species included, and have the same instinctive jealousy or aversion +towards them as against the nymphs of queens properly so denominated. + + + + +LETTER VI. + +_ON THE COMBATS OF QUEENS: THE MASSACRE OF THE MALES: AND WHAT SUCCEEDS +IN A HIVE WHERE A STRANGER QUEEN IS SUBSTITUTED FOR THE NATURAL ONE._ + + +M. de Reaumur had not witnessed every thing relative to bees when he +composed his history of these industrious animals. Several observers, +and those of Lusaçe in particular, have discovered many important facts +that escaped him; and I, in my turn, have made various observations of +which he had no suspicion: at the same time, and this is a very +remarkable circumstance, not only has all that he expressly declares he +saw been verified by succeeding naturalists, but all his conjectures are +found just. The German naturalists, Schirach, Hattorf, and Riems +sometimes contradict him, indeed, in their memoirs; but I can maintain +that, while combating the opinion of M. de Reaumur, it is they who are +almost always wrong; of which several instances might be adduced. + +What I shall now proceed to say will give me an opportunity of detailing +some interesting facts. + +It was observed by M. de Reaumur, that when any supernumerary queen is +either produced in a hive, or comes into it, one of the two soon +perishes. He has not actually witnessed the combat in which she falls, +but he conjectures there is a mutual attack, and that the empire remains +with the strongest or the most fortunate. M. Schirach, on the other +hand, and, after him, M. Riems, thinks that the working bees assail the +stranger, and sting her to death. I cannot comprehend by what means they +have been able to make this observation: as they used very thick hives +only, with several rows of combs, they could at most but observe the +commencement of hostilities. While the combat lasts, the bees move with +great rapidity; they fly on all sides; and, gliding between the combs, +conceal their motions from the observer. For my part, though using the +most favourable hives, I have never seen a combat between the queens and +workers, but I have very often beheld one between the queens themselves. + +In one of my hives in particular, there were five or six royal cells, +each including a nymph. The eldest first underwent its transformation. +Scarcely did ten minutes elapse from the time of this young queen +leaving her cradle, when she visited the other royal cells still close. +She furiously attacked the nearest; and, by dint of labour, succeeded +in opening the top: we saw her tearing the silk of the coccoon with her +teeth; but her efforts were probably inadequate to the object, for she +abandoned this end of the cell, and began at the other, where she +accomplished a larger aperture. When it was sufficiently enlarged, she +endeavoured to introduce her belly, and made many exertions until she +succeeded in giving her rival a deadly wound with her sting. Then having +left the cell, all the bees that had hitherto been spectators of her +labour, began to increase the opening, and drew out the dead body of a +queen scarcely come from its envelope of a nymph. + +Meanwhile, the victorious young queen attacked another royal cell, but +did not endeavour to introduce her extremity into it. There was only a +royal nymph, and no queen, come to maturity, as in the first cell. In +all probability, nymphs of queens inspire their rivals with less +animosity; still they do not escape destruction: because, whenever a +royal cell has been opened before the proper time, the bees extract the +contents in whatever form they may be, whether worm, nymph, or queen. +Lastly, the young queen attacked the third cell, but could not succeed +in penetrating it. She laboured languidly, and appeared as if exhausted +by her first exertions. As we now required queens for some particular +experiments, we resolved to remove the other royal cells, yet in safety, +to secure them from her fury. + +After this observation, we wished to see what ensued on two queens +leaving their cells at the same time, and in what manner one perished. I +find an observation on this head in my Journal, 15. May 1790. + +In one of our thinnest hives, two queens left their cells almost at the +same moment. Whenever they observed each other, they rushed together, +apparently with great fury, and were in such a position that the antennæ +of each was seized by the teeth of the other: the head, breast, and +belly of the one were exposed to the head, breast, and belly of the +other: the extremity of their bodies were curved; they were reciprocally +pierced with the stings; and both fell dead at the same instant. But it +seems as if nature has not ordained that both combatants should perish +in the duel; but rather that, when finding themselves in the situation +described, namely, opposite, and belly to belly, they fly at that moment +with the utmost precipitation. Thus, when these two rivals felt the +extremities about to meet, they disengaged themselves, and each fled +away. You will observe, Sir, that I have repeated this observation very +often, so that it leaves no room for doubt: and I think that we may here +penetrate the intention of nature. + +There ought to be none but one queen in a hive: therefore it is +necessary, if by chance a second is either produced or comes into the +hive, that one of the two must be destroyed. This cannot be committed to +the working bees, because, in a republic composed of so many +individuals, an equal consent cannot be supposed always to exist; it +might frequently happen that one group of bees destroyed one of the +queens, while a second would massacre the other; and the hive thus be +deprived of queens. Therefore it was necessary that the queens +themselves should be entrusted with the destruction of their rivals: but +as, in these combats, nature demands but a single victim, she has wisely +arranged that, at the moment when, from their position, the two +combatants might lose their lives, both feel so great an alarm, that +they think only of flight, and not of using their stings. + +I am well aware of the hazard of error in minute researches into the +causes of the most trifling facts. But here the object and the means +seem so plain, that I have ventured to advance my conjectures. You will +judge better than I can, whether they are well founded.--Let me now +return from this digression. + +A few minutes after the two queens separated, their terror ceased, and +they again began to seek each other. Immediately on coming in sight, +they rushed together, seized one another, and resumed exactly their +former position. The result of this encounter was the same. When their +bellies approached, they hastily disengaged themselves, and fled with +precipitation. During all this time, the workers seemed in great +agitation; and the tumult appeared to increase when the adversaries +separated. Two different times, we observed them stop the flight of the +queens, seize their limbs, and retain them prisoners above a minute. At +last, the queen, which was either the strongest or the most enraged, +darted on her rival at a moment when unperceived, and with her teeth +caught the origin of the wing; then rising above her, brought the +extremity of her own body under the belly of the other; and, by this +means, easily pierced her with the sting. Then she withdrew her sting +after losing hold of the wing. The vanquished queen fell down, dragged +herself languidly along, and, her strength failing, she soon expired. + +This observation proved that virgin queens engage in single combats; but +we wished to discover whether those fecundated, and mothers, had the +same animosity. + +On the 22. of July, we selected a flat hive, containing a very fertile +queen: and being curious to learn whether, as virgin queens, she would +destroy the royal cells, three were introduced into the middle of the +comb. Whenever she observed this, _she_ sprung forward on the whole, and +pierced them towards the bottom; nor did she desist until the included +nymphs were exposed. The workers which had hitherto been spectators of +this destruction, now came to carry the nymphs away. They greedily +devoured the food remaining at the bottom of the cells, and also sucked +the fluid from the abdomen of the nymphs: and then terminated with +destroying the cells from which they had been drawn. + +In the next place, we introduced a very fertile queen into this hive; +after painting the thorax to distinguish her from the reigning queen. A +circle of bees quickly formed around the stranger, but their intention +was not to caress and receive her well; for they insensibly accumulated +so much, and surrounded her so closely, that in scarcely a minute she +lost her liberty and became a prisoner. It is a remarkable circumstance, +that other workers at the same time collected round the reigning queen +and restrained all her motions; we instantly saw her confined like the +stranger. Perhaps it may be said, the bees anticipated the combat in +which these queens were about to engage, and were impatient to behold +the issue of it, for they retained their prisoners only when they +appeared to withdraw from each other; and if one less restrained seemed +desirous of approaching her rival, all the bees forming the clusters +gave way to allow her full liberty for the attack; then if the queens +testified a disposition to fly, they returned to enclose them. + +We have repeatedly witnessed this fact, but it presents so new and +singular a characteristic in the policy of bees, that it must be seen +again a thousand times before any positive assertion can be made on the +subject. I would therefore recommend that naturalists should attentively +examine the combat of queens, and particularly ascertain what part is +taken by the workers. Is their object to accelerate the combat? Do they +by any secret means excite the fury of the combatants? Whence does it +happen that accustomed to bestow every care on their queen, in certain +circumstances, they oppose her preparations to avoid impending danger? + +A long series of observations are necessary to solve these problems. It +is an immense field for experiment, which will afford infinitely curious +results. I intreat you to pardon my frequent digressions. The subject is +deeply philosophical, genius such as your's is required to treat it +properly; and I shall now be satisfied with proceeding in the +description of the combat. + +The cluster of bees that surrounded the reigning queen having allowed +her some freedom, she seemed to advance towards that part of the comb +where her rival stood; then, all the bees receded before her, the +multitude of workers, separating the two adversaries, gradually +dispersed, until only two remained; these also removed, and allowed the +queens to come in sight. At this moment, the reigning queen rushed on +the stranger, with her teeth seized her near the origin of the wing, and +succeeded in fixing her against the comb without any possibility of +motion or resistance. Next curving her body, she pierced this unhappy +victim of our curiosity with a mortal wound. + +In the last place, to exhaust every combination, we had still to examine +whether a combat would ensue between two queens, one impregnated, and +the other a virgin; and what circumstances attended it. + +On the 18. of September, we introduced a very fruitful queen into a +glass hive, already containing a virgin queen, and put her on the +opposite side of the comb, that we might have time to see how the +workers would receive her. She was immediately surrounded, but they +confined her only a moment. Being oppressed with the necessity of +laying, she dropped some eggs; however, we could not discover what +became of them; certainly the bees did not convey them to the cells, +for, on inspection, we found none there. The group surrounding this +queen having dispersed a little, she advanced towards the edge of the +comb, and soon approached very near the virgin queen. When in sight, +they rushed together; the virgin queen got on the back of the other, and +gave her several stings in the belly, but, having aimed at the scaly +part, they did not injure her, and the combatants separated. In a few +minutes, they returned to the charge; but this time the impregnated +queen mounted on her rival; however, she sought in vain to pierce her, +for the sting did not enter; the virgin queen then disengaged herself +and fled; she also succeeded in escaping another attack, where her +adversary had the advantage of position. These rivals appeared nearly of +equal strength; and it was difficult to foresee to which side victory +would incline, until at last, by a successful exertion, the virgin queen +mortally wounded the stranger, and she expired in a moment. The sting +had penetrated so far that the victorious queen was unable to extract +it, and she was overthrown by the fall of her enemy. She made great +exertions to disengage the sting: but could succeed by no other means +than turning on the extremity of the belly, as on a pivot. Probably the +barbs of the sting fell by this motion, and, closing like a spiral +around the stem, came more easily from the wound. + +These observations, Sir, I think will satisfy you, respecting the +conjecture of our celebrated Reaumur. It is certain, that if several +queens are introduced into a hive, one alone will preserve the empire; +that the others will perish from her attacks; and that the workers will +at no time attempt to employ their stings against the stranger queen. I +can conceive what has misled M. Riems and Schirach; but it is necessary +for explaining it that I should relate a new feature in the policy of +bees, at considerable length. + +In the natural state of hives, several queens from different royal +cells, may sometimes exist at the same moment, and they will remain +either until formation of a swarm or a combat among them decides to +which the throne shall appertain. But excepting this case, there never +can be supernumerary queens; and if an observer wishes to introduce one, +he can accomplish it only by force, that is by opening the hive. In a +word, no queen can insinuate herself into a hive in a natural state, +for the following reasons. + +Bees preserve a sufficient guard, day and night, at the entrance of +their habitation. These vigilant centinels examine whatever is +presented; and, as if distrusting their eyes, they touch with the +antennæ every individual endeavouring to penetrate the hive, and also +the various substances put within their reach; which affords us an +opportunity of observing that the antennæ are certainly the organs of +feeling. If a stranger queen appears, she is instantly seized by the +bees on guard, which prevent her entry by laying hold of her legs or +wings with their teeth, and crowd so closely around her, that she cannot +move. Other bees, from the interior of the hive, gradually come to their +assistance, and confine her still more narrowly, all having their heads +towards the centre where the queen is inclosed; and they remain with +such evident anxiety, eagerness, and attention, that the cluster they +form may be carried about for some time, without their being sensible +of it. A stranger queen, so closely confined and hemmed in, cannot +possibly penetrate the hive. If the bees retain her too long imprisoned, +she perishes. Her death probably ensues from hunger, or the privation of +air; it is undoubted, at least, that she is never stung. We never saw +the bees direct their stings against her, except a single time, and then +it was owing to ourselves. We endeavoured, from compassion for a queen's +situation, to remove her from the center of a cluster; the bees became +enraged; and, in darting out their stings, some struck the queen, and +killed her. It is so certain that the stings were not purposely directed +against her, that several of the workers were themselves killed; and +surely they could not intend destroying one another. Had we not +interfered, they would have been content with confining the queen, and +would not have massacred her. + +It was in similar circumstances that M. Riems saw the workers anxiously +pursue a queen. He thought they designed to sting her, and thence +concluded, that the office of the common bees is to kill supernumerary +queens. You have quoted his observations in the _Contemplation de la +Nature, part II, chap. 27, note 7_. But you are sensible, Sir, from +these details, that he has been mistaken. He did not know the attention +that bees bestow on what passes at the entrance of their hive, and he +was entirely ignorant of the means they take to prevent supernumerary +queens from penetrating it. + + * * * * * + +After ascertaining that the workers in no situation sting the +supernumerary queens, we were curious to learn how a stranger queen +would be received in a hive wanting a reigning one. To elucidate this +matter, we made numerous experiments, the detail of which would protract +this letter too much, therefore I shall relate only the principal +results. + +Bees do not immediately observe the removal of their queen; their +labours are uninterrupted; they watch over the young, and perform all +their ordinary occupations. But, in a few hours, agitation ensues; all +appears a scene of tumult in the hive. A singular humming is heard; the +bees desert their young; and rush over the surface of the combs with a +delirious impetuosity. Then they discover their queen is no longer among +them. But how do they become sensible of it? How do the bees on the +surface of the comb discover that the queen is not on the next comb? In +treating of another characteristic of these animals, you have yourself, +Sir, proposed the same question; I am incapable of answering it indeed, +but I have collected some facts, that may perhaps facilitate the +elucidation of this mystery. + +I cannot doubt that the agitation arises from the workers having lost +their queen; for on restoring her, tranquillity is instantly regained +among them; and, what is very singular, they _recognise_ her: you must +interpret this expression strictly. Substitution of another queen is not +attended with the same effect, if she is introduced into the hive within +the first twelve hours after removal of the reigning one. Here the +agitation continues; and the bees treat the stranger the same as when +the presence of their own leaves them nothing to desire. They surround, +seize, and keep her captive, a very long time, in an impenetrable +cluster; and she commonly dies either from hunger or privation of air. + +If eighteen hours elapse before substitution of a stranger queen for the +native one removed, she is at first treated in the same manner, but the +bees leave her sooner; nor is the surrounding cluster so close; they +gradually disperse; and the queen is at last liberated. She moves +languidly; and sometimes expires in a few minutes. However some queens +have escaped in good health from an imprisonment of seventeen hours; and +ended with reigning in the hives where they had originally been ill +received. + +If, before substituting the stranger queen, twenty-four hours elapse, +she will be well received, and reign from the moment of her introduction +into the hive. Here I speak of the good reception given to a queen after +an interregnum of twenty-four hours. But as this word reception is very +indefinite, it is proper to enter into some detail for explaining the +exact sense in which I use it. On the 15. of August, I introduced a +fertile queen, eleven months old, into a glass hive. The bees were +twenty-four hours deprived of their queen, and had already begun the +construction of twelve royal-cells, such as described in the preceding +chapter. Immediately on placing this female stranger on the comb, the +workers near her touched her with their antennæ, and, passing their +trunks over every part of her body, they gave her honey. Then these gave +place to others that treated her exactly in the same manner. All +vibrated their wings at once, and ranged themselves in a circle around +their sovereign. Hence resulted a kind of agitation which gradually +communicated to the workers situated on the same surface of the comb, +and induced them to come and reconnoitre, in their turn, what was going +on. They soon arrived; and, having broke through the circle formed by +the first, approached the queen, touched her with the antennæ, and gave +her honey. After this little ceremony they retired; and, placing +themselves behind the others, enlarged the circle. There they vibrated +their wings, and buzzed without tumult or disorder, and as if +experiencing some very agreeable sensation. The queen had not yet moved +from the place where I had put her, but in a quarter of an hour she +began to move. The bees, far from opposing her, opened the circle at +that part to which she turned, followed her, and formed a guard around. +She was oppressed with the necessity of laying, and dropped eggs. +Finally, after four hours abode, she began to deposit male eggs in the +cells she met. + +While these events passed on the surface of the comb where the queen +stood, all was quiet on the other side. Here the workers were apparently +ignorant of a queen's arrival in the hive. They laboured with great +activity at the royal cells, as if ignorant that they no longer stood in +need of them: they watched over the royal worms, supplied them with +jelly and the like. But the queen having at length come to this side, +she was received with the same respect that she had experienced from +their companions on the other side of the comb. They encompassed her; +gave her honey; and touched her with their antennæ: and what proved +better that they treated her as a mother, was their immediately +desisting from work at the royal cells; they removed the worms, and +devoured the food collected around them. From this moment the queen was +recognised by all her people, and conducted herself in this new +habitation as if it had been her native hive. + +These particulars will give a just idea of the manner that bees receive +a stranger queen; when they have time to forget their own, she is +treated exactly as if she was their natural one, except that there is +perhaps at first greater interest testified in her, or more conspicuous +demonstrations of it. I am sensible of the impropriety of these +expressions, but M. de Reaumur in some respect authorises them. He does +not scruple to say, that bees pay _attention_, _homage_, and _respect_, +to their queen, and from his example the like expressions have escaped +most authors that treat on bees. + +Twenty-four or thirty hours absence is sufficient to make them forget +their first queen, but I can hazard no conjecture on the cause. + + * * * * * + +Before terminating this letter, which is full of combats and disastrous +scenes, I should, perhaps, give you an account of some more pleasing +and interesting facts relative to their industry. However, to avoid +returning to duels and massacres, I shall here subjoin my observations +on the massacre of the males. + +You will remember, Sir, it is agreed by all observers, that at a certain +period of the year, the workers kill and expel the drones. M. de Reaumur +speaks of these executions as a horrible massacre. He does not expressly +affirm, indeed, that he has himself witnessed it, but what we have seen +corresponds so well with his account, that there can be no doubt he has +beheld the peculiarities of the massacre. + +It is usually in the months of July and August, that the bees free +themselves of the males. Then they are drove away and pursued to the +inmost parts of the hive, where they collect in numbers; and as at the +same time we find many dead drones on the ground before the hives, it is +indubitable that after being expelled, the bees sting them to death. +Yet on the surface of the comb, we do not see the sting used against +them; there the bees are content to pursue and drive them away. You +observe this, Sir, yourself, in the new notes added to _la Contemplation +de la Nature_; and you seem disposed to think, that the drones forced to +retire to the extremity of the hive, perish from hunger. Your conjecture +was extremely probable. Still it was possible the carnage might take +place in the bottom of the hive, and had been unobserved, because that +part is dark, and escapes the observer's eye. + +To appreciate the justice of this suspicion, we thought of making the +support of the hive of glass, and of placing ourselves below to see what +passed in the scene of action. Therefore, a glass table was constructed, +on which were put six hives with swarms of the same year; and, lying +under it, we endeavoured to discover how the drones were destroyed. The +invention succeeded to admiration. On the 4 of July, we saw the workers +actually massacre the males, in the whole six swarms, at the same hour, +and with the same peculiarities. + +The glass table was covered with bees full of animation, which flew upon +the drones, as they came from the bottom of the hive; seized them by the +antennæ, the limbs, and the wings, and after having dragged them about, +or, so to speak, after quartering them, they killed them by repeated +stings directed between the rings of the belly. The moment that this +formidable weapon reached them, was the last of their existence; they +stretched their wings, and expired. At the same time, as if the workers +did not consider them as dead as they appeared to us, they still stuck +the sting so deep, that it could hardly be withdrawn, and these bees +were obliged to turn upon themselves before the stings could be +disengaged. + +Next day, having resumed our former position, we witnessed new scenes of +carnage. During three hours, the bees furiously destroyed the males. +They had massacred all their own on the preceding evening, but now +attacked those which, driven from the neighbouring hives, had taken +refuge amongst them. We saw them also tear some remaining nymphs from +the combs; they greedily sucked all the fluid from the abdomen, and then +carried them away. The following days no drones remained in the hives. + +These two observations seem to me decisive. It is incontestible that +nature has charged the workers with the destruction of the males at +certain seasons of the year. But what means does she use to excite their +fury against them? This is a question that I cannot pretend to answer. +However, an observation I have made may one day lead to solution of the +problem. The males are never destroyed in hives deprived of queens, on +the contrary, while a savage massacre prevails in other places, they +there find an asylum. They are tolerated and fed, and many are seen even +in the middle of January. They are also preserved in hives, which, +without a queen properly so called, have some individuals of that +species that lay the eggs of males, and in those whose half fecundated +queens, if I may use the expression, propagate only drones. Therefore, +the massacre takes place but in hives where the queens are completely +fertile, and it never begins until the season of swarming is past. + + _PREGNY, 28 August 1791._ + + + + +LETTER VII. + +_SEQUEL OF EXPERIMENTS ON THE RECEPTION OF A STRANGER QUEEN. M. DE +REAUMUR'S OBSERVATIONS ON THE SUBJECT._ + + +I have frequently testified my admiration of M. de Reaumur's +observations on bees. I feel a sensible pleasure in acknowledging that +if I have made any progress in the art of observation, I am indebted for +it to profound study of the works of this naturalist. In general his +authority has such weight, that I can scarcely trust my own experiments +when the results are different from his. Likewise, on finding myself in +opposition to the _historian of bees_, I repeat my experiments. I vary +the mode of conducting them; I examine with the utmost caution all the +circumstances that might mislead me, and never are my labours +interrupted before acquiring the moral certainty of avoiding error. With +the aid of these precautions, I have discovered the justice of M. de +Reaumur's suggestions, and I have a thousand times seen, if certain +experiments seemed to combat them, it was from incorrectness of +execution. Yet I must except some facts where my results have constantly +been different from his. Those respecting the reception of a stranger +queen substituted for the natural one, are of the number. + +If, after removing the natural queen, a stranger is immediately +substituted, the usurper is ill received. I never could succeed in +making them adopt her, but by allowing an interval of twenty or +twenty-four hours to elapse. Then they seemed to have forgot their own +queen; and respectfully received any female put in her place. M. de +Reaumur, on the contrary, asserts, that should the original queen be +removed, and another presented, this new one will be perfectly well +received from the beginning. As evidence of this assertion, he gives the +detail of an experiment which must be read in his work, for I shall here +give only an extract of it{L}. He induced four or five hundred bees to +leave their native hive and enter a glass box, containing a small piece +of comb towards the top. At first they were in great agitation; and, to +pacify or console them, he presented a new queen. From this moment, the +tumult ceased, and the stranger queen was received with all respect. + +I do not dispute the truth of this experiment; but, in my opinion, it +does not warrant the conclusion that M. de Reaumur deduces from it. His +apparatus removed the bees too much from their natural condition, to +allow him to judge of their instinct and dispositions. In other +situations, he has himself observed, that these animals, reduced to +small numbers, lost their industry and activity, and feebly continued +their ordinary labours. Thus their instinct is affected by every +operation that too much diminishes their number. To render such an +experiment truly conclusive, it must be made in a populous hive; and on +removing the native queen, a stranger must immediately be substituted in +her place. Had this been done, I am fully persuaded, that M. de Reaumur +would have seen the bees imprison the usurper, confine her at least +twelve or fifteen hours among them, and frequently suffocate her: nor +would he have witnessed any favourable reception before an interval of +twenty-four hours after removal of the original queen. No variation has +occurred in my experiments regarding this fact. Their number, and the +attention bestowed on them, make me presume they merit your confidence. + +M. de Reaumur, in another passage of the same Memoir, affirms, that +_bees, which have a queen they are satisfied with, are nevertheless +disposed to give the best possible reception to any female that seeks +refuge among them_. In the preceding letter, I have related my +experiments on this head: their success has been very different from +that of M. de Reaumur's. I have proved that the workers never employ +their stings against the queen; but this cannot be called the welcome +reception of a stranger. They retain her within their ranks, and seem to +allow her liberty only when she prepares to combat the reigning queen. +This observation cannot be made except in the thinnest hives. Those used +by M. de Reaumur had always two parallel combs at least, which must +have prevented him from observing some very important circumstances that +influence the conduct of workers when supplied with several females. The +first circles formed around a stranger queen he has taken for caresses; +and, from the little that this queen could advance between the combs, it +must have been impossible for him to observe that the circles, which +always continued contracting, ended in restraint of the females there +inclosed. Had he used thinner hives, he would have discovered that what +he supposed indication of a favourable reception was the prelude of +actual imprisonment. + +I feel reluctant to assert that M. de Reaumur was deceived. Yet I cannot +admit that, on certain occasions, bees tolerate a plurality of females +in their hives. The experiment on which this affirmation rests will not +be considered decisive. In the month of December, he introduced a +stranger queen into a glass hive, in his cabinet, and confined her +there. The bees had no opportunity of going out. This stranger was well +received; her presence awakened the workers from their lethargic state, +into which they did not relapse; she excited no carnage; the number of +dead bees on the board of the hive did not sensibly increase; and no +dead queens were found. + +Before concluding any thing favourable to the plurality of queens, it +was necessary to ascertain whether the native queen was living when the +new one was introduced into the hive: however the author neglected this; +and it is very probable the hive had lost its queen, since the bees were +languid, and the presence of a stranger restored their activity. + +I trust, Sir, that you will pardon this slight criticism. Far from +industriously seeking faults in our celebrated Reaumur, I derive the +greatest pleasure when my observations coincide with his, and still +more, when my experiments justify his conjectures. But I think it +proper to point out those cases where the imperfections of his hives +have led him into error, and to explain from what causes I have not seen +certain facts in the same manner he did. I feel particular anxiety to +merit your confidence, and I am aware that the greatest exertions are +necessary, when I have to combat the historian of bees. I confide in +your judgment; and pray you to be assured of my respect. + + _PREGNY, 30. August 1791._ + + +FOOTNOTES: + +{L} Edit. 4to, Tom. V. p. 258. + + + + +LETTER VIII. + +_IS THE QUEEN OVIPAROUS? WHAT INFLUENCE HAS THE SIZE OF THE CELLS, WHERE +THE EGGS ARE DEPOSITED, ON THE BEES PRODUCED?--RESEARCHES ON THE MODE OF +SPINNING THE COCCOONS._ + + +In this letter I shall collect some isolated observations relative to +various points in the history of bees, concerning which you wished me to +engage. + +You desired me to investigate whether the queen is really _oviparous_. +M. de Reaumur leaves this question undecided. He observes, that he has +never seen the worm hatched; and he only asserts that worms are found in +those cells where eggs have been deposited three days preceding. If we +attempt to catch the moment when the worm leaves the egg, we must extend +our observations beyond the interior of the hive; for there the +continual motion of the bees obscures what passes at the bottom of +cells. The egg must be taken out, presented to the microscope, and every +change attentively watched. One other precaution is essential. As a +certain degree of heat is requisite to hatch the worms, should the eggs +be too soon deprived of it they wither and perish. The sole method of +succeeding in seeing the worm come out, consists in watching the queen +while she lays, in marking the egg so as to be recognised, and removing +it from the hive to the microscope only an hour or two before the three +days elapse. The worm will certainly be hatched, provided it has been +exposed as long as possible to the full degree of heat. Such is the +course I have pursued; and the following are the results obtained. + +In the month of August, we removed several cells containing eggs that +had been three days deposited: we cut off the top of the cell, and put +the pyramidal bottom, where the egg was fixed, on a glass slider. Slight +motions were soon perceptible in the eggs. At first, we could observe no +external organization: the worm was entirely concealed from us by its +pellicle. We then prepared to examine the egg with a powerful magnifier; +however, during the interval, the worm burst its surrounding membrane, +and cast off part of the envelope, which was torn and ragged on +different parts of the body, and more evidently so towards the last +rings. The worm alternately curved and stretched itself, with very +lively action. Twenty minutes were occupied in casting off the spoil; +when this exertion ceased: the worm lay down, curved, and seemed to take +that rest which it required. An egg laid in a worker's cell produced +this animal, which would have become a worker itself. + +We next directed our attention to the moment when a male worm would be +hatched. An egg was exposed to the sun on a glass slider; and, with a +good magnifier, nine rings of the worm were perceptible within the +transparent pellicle. This membrane was still entire, and the worm +perfectly motionless. The two longitudinal lines of tracheæ were visible +on the surface, and many ramifications. We never lost sight of the egg a +single instant, and now succeeded in observing the first motions of the +worm. The thick end alternately straightened and curved, and almost +reached the part where the sharp extremity was fixed. These exertions +burst the membrane, first on the upper part, towards the head, then on +the back, and afterwards on all the rest successively. The ragged +pellicle remained in folds on different parts of the body, and then fell +off. Thus it is beyond dispute, that the queen is oviparous. + +Some observers affirm, that the workers attend to the eggs before the +worms are hatched; and it is certain that, at whatever time a hive is +examined, we always see some workers with the head and thorax inserted +into cells containing eggs, and remaining motionless several minutes in +this position. It is impossible to discover what they do, for the +interior of the cell is concealed by their bodies; but it is very easily +ascertained that, in this attitude, they are doing nothing to the eggs. + +If, at the moment the queen lays, her eggs are put into a grated box, +and deposited in a strange hive, where there is the necessary degree of +heat, the worms come out at the usual time, just as if they had been +left in the cells. Thus no extraordinary aid or attention is required +for their exclusion. + +When the workers penetrate the cells, and remain fifteen or twenty +minutes motionless, I have reason to believe, it is only to repose from +their labours. My observations on the subject seem correct. You know, +Sir, that a kind of irregular shaped cells, are frequently constructed +on the panes of the hive. These, being glass on one side, are +exceedingly convenient to the observer, since all that passes within is +exposed. I have often seen bees enter these cells when nothing could +attract them. The cells contained neither eggs nor honey, nor did they +need further completion. Therefore the workers repaired thither only to +enjoy some moments of repose. Indeed, they were fifteen or twenty +minutes so perfectly motionless, that had not the dilatation of the +rings shewed their respiration, we might have concluded them dead. The +queen also sometimes penetrates the large cells of the males, and +continues very long motionless in them. Her position prevents the bees +from paying their full homage to her, yet even then the workers do not +fail to form a circle around her, and brush the part of her belly that +remains exposed. + +The drones do not enter the cells while reposing, but cluster together +on the combs; and sometimes retain this position eighteen or twenty +hours without the slightest motion. + +As it is important, in many experiments, to know the exact time that the +three species of bees exist before assuming their ultimate form, I shall +here subjoin my own observations on the point. + +The worm of workers passes three days in the egg, five in the vermicular +state, and then the bees close up its cell with a wax covering. The worm +now begins spinning its coccoon, in which operation thirty-six hours are +consumed. In three days, it changes to a nymph, and passes six days in +this form. It is only on the twentieth day of its existence, counting +from the moment the egg is laid, that it attains the fly state. + +The royal worm also passes three days in the egg, and is five a worm; +the bees then close its cell; and it immediately begins spinning the +coccoon, which occupies twenty-four hours. The tenth and eleventh day it +remains in complete repose, and even sixteen hours of the twelfth. Then +the transformation to a nymph takes place, in which state four days and +a third are passed. Thus it is not before the sixteenth day that the +perfect state of queen is attained. + +The male worm passes three days in the egg, six and a half as a worm, +and metamorphoses into a fly on the twenty-fourth day after the egg is +laid. + +Though the larvæ of bees are apodal, they are not condemned to absolute +immobility in their cells; for they can move by a spiral motion. During +the first three days, this motion is so slow as scarcely to be +perceptible, but it afterwards becomes more evident. I have then +observed them perform two complete revolutions in an hour and three +quarters. When the period of transformation arrives, they are only two +lines from the orifice of the cells. As their position is constantly the +same, bent in an arc, those in the workers' and drones' cells are +perpendicular to the horizon, while those in the royal cells lie +horizontally. It might be thought, that the difference of position has +much influence on the increment of the different larvæ; yet it has none. +By reversing combs containing common cells full of brood, I have put the +worms in a horizontal position; but they were not injured. I have also +turned the royal cells, so that the worms came into a horizontal +direction; however their increment was neither slower nor less perfect. + + * * * * * + +I have been much surprised at the mode of bees spinning their coccoons, +and I have witnessed many new and interesting facts. The worms both of +workers and males fabricate _complete_ coccoons in their cells; that +is, close at both ends, and surrounding the whole body. The royal larvæ, +on the other hand, spin imperfect coccoons, open behind, and enveloping +only the head, thorax, and first ring of the abdomen. The discovery of +this difference, which at first may seem trifling, has given me extreme +pleasure, for it evidently demonstrates the admirable art with which +nature connects the various characteristics in the industry of bees. + +You will remember, Sir, the evidence I gave you of the mutual aversion +of queens, of the combats in which they engage, and the animosity that +leads them to destroy one another. Of several royal nymphs in a hive, +the first transformed attacks the rest, and stings them to death. But +were these nymphs enveloped in a complete coccoon, she could not +accomplish it. Why? because the silk is of so close a texture, the sting +could not penetrate, or if it did, the barbs would be retained by the +meshes of the coccoon, and the queen, unable to retract it, would +become the victim of her own fury. Thus, that the queen might destroy +her rivals, it was necessary the last rings of the body should remain +uncovered, therefore the royal nymphs must only form imperfect coccoons. +You will observe, that the last rings alone should be exposed, for the +sting can penetrate no other part: the head and thorax are protected by +connected shelly plates which it cannot pierce. + +Hitherto, philosophers have claimed our admiration of nature in her care +of preserving and multiplying the species. But from the facts I relate, +we must admire her precautions in exposing certain individuals to a +mortal danger. + +The detail on which I have just entered clearly indicates the final +cause of the opening left by the royal worms in their coccoons; but it +does not shew whether it is in consequence of a particular instinct that +they leave this opening, or whether the wideness of their cells prevents +them from stretching the thread up to the top. This question interested +me very much; the only method of deciding it was to observe the worms +while spinning, which cannot be done in their opaque cells. It then +occurred to me to dislodge them from their own habitations, and +introduce them into glass tubes, blown in exact imitation of the +different kind of cells. The most difficult part of the operation +consisted in extracting worms and introducing them here; but my +assistant accomplished it with much address. He opened several sealed +royal cells, where we knew the larvæ were about to begin their coccoons, +and, taking them gently out, introduced one into each of my glass cells +without the smallest injury. + +They soon prepared to work; and commenced by stretching the anterior +part of the body in a straight line, while the other was bent in a +curve. This formed a curve of which the longitudinal sides of the cells +were tangents, and afforded two points of support. The head was next +conducted to the different parts of the cell which it could reach, and +it carpeted the surface with a thick bed of silk. We remarked that the +threads were not carried from one side to another, and that this would +have been impracticable, for the worms being obliged to support +themselves, and to keep the posterior rings curved, the free and +moveable part of the body was not long enough for the mouth to reach the +sides diametrically opposite, and fix the threads to them. You will +remember, Sir, that the royal cells are of a pyramidal form, with a wide +base, and a long contracted top. These cells hang perpendicularly in the +hive, the point downwards, from which position the royal worm can be +supported in the cell, only when the curvature of the posterior part +forms two points of support; and that it cannot obtain this support +without resting on the lower part, or towards the extremity. Therefore +if it attempted to stretch out and spin towards the wide end of the +cell, it could not reach both sides from being too distant. One part +would be touched by its extremity, the other by its back, and it would +consequently tumble down. I have particularly ascertained the fact in +glass cells that were too large, and of which the diameter was greater +towards the point than is usual in cells; there they were unable to +support themselves. + +These first experiments obviated the suspicion of any particular +instinct in the royal worms. They proved, if the worms spun incomplete +coccoons, it was because they were forced to do so by the figure of +their cells. However, I wished to have evidence still more direct. I put +them into cylindrical glass cells, or portions of glass tubes resembling +common cells, and I had the satisfaction of seeing them spin complete +coccoons, as the worms of workers do. Lastly, I put common worms in very +wide cells, and they left the coccoon open. Thus it is demonstrated, +that the royal worms, and those of workers, have the same instinct and +the same industry, or in other words, when situated in the same +circumstances, the course they follow is the same. I may here add, that +the royal worms artificially lodged in cells, where they can spin +complete coccoons, undergo all their metamorphoses equally well. Thus +the necessity imposed on them by nature, of having the coccoons open, is +not necessary for their increment; nor has it any other object than that +of exposing them to the certainty of perishing by the wounds of their +natural enemy; an observation new and truly singular. + + * * * * * + +I ought to relate my experiments on the influence that the size of the +cells has on bees. It is to you, Sir, that I am indebted for suggesting +them. + +As we sometimes find males smaller than they ought to be, and also +queens more diminutive than usual, it was desirable to obtain a general +explanation, to what degree the cells, where bees pass the first period +of their existence, influence their size. With this view, you have +advised me to remove all the combs composed of common cells, and to +leave those consisting of large cells only. It was evident if the common +eggs which the queen would lay in these large cells produced workers of +larger size, we were bound to conclude that the size of the cells had a +sensible influence on the size of the bees. The first time I made this +experiment, it did not succeed, because weevils lodged in the hive +discouraged the bees. But I repeated it afterwards, and the result was +very remarkable. + +I removed the whole comb, consisting of common cells, from one of my +best glass hives, and left that composed of males' cells alone: and to +avoid vacuities, I supplied others of the same kind. This was in June, +the season most favourable to bees. I expected that the bees would +quickly have repaired the ravages produced by this operation in their +dwelling; that they would labour at the breaches, and unite the new +combs to the old. But I was very much surprised to see that they did not +begin to work. Expecting they would resume their activity, I continued +observing them several days; however, my hopes were disappointed. Their +homage to the queen was not interrupted indeed; but except in this, +their conduct to the queen was quite different from what it usually is; +they clustered on the combs without exciting any sensible heat. A +thermometer among them rose only to 81°, though standing at 77° in the +open air. In a word, they appeared in a state of the greatest +despondency. + +The queen herself, though very fertile, and though she must have been +oppressed by her eggs, hesitated long before depositing them in the +large cells; she chose rather to drop them at random than lay in cells +unsuitable. However, on the second day, we found six that had been +deposited there with all regularity. The worms were hatched three days +afterwards, and then we began to study their history. Though the bees +provided them with food, they did not carefully attend to it; yet I was +in hopes they might be reared. I was again disappointed; for next +morning all the worms had disappeared, and their cells were left empty. +Profound silence reigned in the hive; few bees left it, and these +returned without pellets of wax on the limbs; all was cold and +inanimate. To promote a little motion, I thought of supplying the hive +with a comb, composed of large cells, full of male brood of all ages. +The bees, which had twelve days obstinately refused working in wax, did +not unite this comb to their own. However, their industry was awakened +in a way that I had not anticipated. They removed all the brood from +this comb, cleaned out the whole cells, and prepared them for receiving +new eggs. I cannot determine whether they expected the queen to lay, but +it is certain if they did so they were not deceived. From this moment, +she no longer dropped her eggs; but laid such a number in the new comb, +that we found five or six together in the same cell. I then removed all +the combs composed of large cells to substitute small cells in their +place, an operation which restored complete activity among the bees. + +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 larvæ, 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 assistant +substituted 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 substituted 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 larvæ 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 space for that purpose; therefore more would be useless. +Their expansion ought to be no greater in the most spacious 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 larvæ +of workers in the same circumstances. 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 retarded. 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 +spacious 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 assistant took a certain number from the +smallest cells, and deposited them on a quantity 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 sexual 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 assert 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 circumstances 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 passing 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 glass 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 +stalactites, on the edge of the passage 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 antennæ. 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 glass 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 +hitherto 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 assiduity 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 glass 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 passages through the combs, and shaped +liked stalactites. + +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 passed 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 +antennæ, 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 passage 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 participated 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 glass 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 larvæ 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 circumstance, 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 coccoon 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 coccoon. 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 passed within was pretty +perceptible; we could discern that the silk of the coccoon 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 assuming this +attitude, 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, assiduously 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 circumstances, she emitted her sound, assuming 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 coccoon 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. + +The queen this day between twelve and one became extremely agitated. The +royal cells had multiplied very much; she could go no where without +meeting them, and on approaching she was very roughly treated. Then she +fled, but to obtain no better reception. At last, these things agitated +the bees; they precipitately rushed through the outlets of the hive, and +settled on a tree in the garden. It singularly happened that the queen +was herself unable to follow or conduct the swarm. She had attempted to +pass between two royal cells before they were abandoned by the bees +guarding them, and she was so confined and maltreated as to be incapable +of moving. We then removed her into a separate hive prepared for a +particular experiment; the bees, which had clustered on a branch, soon +discovered their queen was not present, and returned of their own accord +to the hive. Such is an account of the second colony of this hive. + +We were extremely solicitous to ascertain what would become of the other +royal cells. Four of the close ones had attained complete maturity, and +the queens would have left them had not the bees prevented it. They were +not open either previous to the agitation of the swarms, or at the +moment of swarming. + +None of the queens were at liberty on the eleventh. The second should +have transformed on the eighth; thus she had been three days confined, a +longer period than the first which formed the swarm. We could not +discover what occasioned the difference in their captivity. + +On the twelfth, the queen was at last liberated, as we found her in the +hive. She had been treated exactly as her predecessor; the bees allowed +her to rest in quiet, when distant from the royal cells, but tormented +her cruelly when she approached them. We watched this queen a long time, +but not aware that she would lead out a colony, we left the hive for a +few hours. Returning at mid-day, we were greatly surprised to find it +almost totally deserted. During our absence, it had thrown a prodigious +swarm, which still clustered on the branch of a neighbouring tree. We +also saw with astonishment the third cell open, and its top connected to +it as by a hinge. In all probability the captive queen, profiting by the +confusion that preceded the swarming, escaped. Thus, there was no doubt +of both queens being in the swarm. We found it so; and removed them, +that the bees might return to the hive, which they did very soon. + +While we were occupied in this operation, the fourth captive queen left +her prison, and the bees found her on returning. At first they were very +much agitated, but calmed towards the evening, and resumed their wonted +labours. They formed a strict guard around the royal cells, and took +great care to remove the queen whenever she attempted to approach. +Eighteen royal cells now remained to be guarded. + +The fifth queen left her cell at ten at night; therefore two queens were +now in the hive. They immediately began fighting, but came to disengage +themselves from each other. However they fought several times during the +night without any thing decisive. Next day, the thirteenth, we witnessed +the death of one, which fell by the wounds of her enemy. This duel was +quite similar to what is said of the combats of queens. + +The victorious queen now presented a very singular spectacle. She +approached a royal cell, and took this moment to utter the sound, and +assume that posture, which strikes the bees motionless. For some +minutes, we conceived, that taking advantage of the dread exhibited by +the workers on guard, she would open it, and destroy the young female; +also she prepared to mount the cell; but in doing so she ceased the +sound, and quitted that attitude which paralyses the bees. The guardians +of the cell instantly took courage; and, by means of tormenting and +biting the queen, drove her away. + +On the fourteenth, the sixth young queen appeared, and the hive threw a +swarm, with all the concomitant disorder before described. The agitation +was so considerable, that a sufficient number of bees did not remain to +guard the royal cells, and several of the imprisoned queens were thus +enabled to make their escape. Three were in the cluster formed by the +swarm, and other three remained in the hive. We removed those that had +led the colony, to force the bees to return. They entered the hive, +resumed their post around the royal cells, and maltreated the queens +when approaching. + +A duel took place in the night of the fifteenth, in which one queen +fell. We found her dead next morning before the hive; but three still +remained, as one had been hatched during night. Next morning we saw a +duel. Both combatants were extremely agitated, either with the desire of +fighting, or the treatment of the bees, when they came near the royal +cells. Their agitation quickly communicated to the rest of the bees, and +at mid-day they departed impetuously with the two females. This was the +fifth swarm that had left the hive between the thirtieth of May and +fifteenth of June. On the sixteenth, a sixth swarm cast, which I shall +give you no account of, as it shewed nothing new. + +Unfortunately we lost this, which was a very strong swarm; the bees flew +out of sight, and could never be found. The hive was now very thinly +inhabited. Only the few bees that had not participated in the general +agitation remained, and those that returned from the fields after the +swarm had departed. The cells were, therefore, slenderly guarded; the +queens escaped from them, and engaged in several combats, until the +throne remained with the most successful. + +Notwithstanding the victories of this queen, she was treated with great +indifference from the sixteenth to the nineteenth, that is, the three +days that she preserved her virginity. At length, having gone to seek +the males, she returned with all the external signs of fecundation, and +was henceforth received with every mark of respect; she laid her first +eggs forty-six hours after fecundation. + +Behold, Sir, a simple and faithful account of my observations on the +formation of swarms. That the narrative might be the more connected, I +have avoided interrupting it by the detail of several particular +experiments which I made at the same time for elucidating various +obscure points of their history. These shall be the subject of future +letters. For, although I have said so much, I hope still to interest +you. + + _PREGNY, 6. September 1791._ + +_P. S._--In revising this letter, I find I have neglected taking notice +of an objection that may embarrass my readers, and which ought to be +answered. + +After the first five swarms had thrown, I had always returned the bees +to the hive: it is not surprising, therefore, that it was continually so +sufficiently stocked that each colony was numerous. But things are +otherwise in the natural state: the bees composing a swarm do not return +to the hive; and it will undoubtedly be asked, What resource enables a +common hive to swarm three or four times without being too much +weakened? + +I cannot lessen the difficulty. I have observed that the agitation, +which precedes the swarming, is often so considerable, that most of the +bees quit the hive, and in that case we cannot well comprehend how, in +three or four days afterwards, it can be in a state to send out another +colony equally strong. + +But remark, in the first place, that the queen leaves a prodigious +quantity of workers' brood, which soon transforms to bees; and in this +way the population sometimes becomes almost as great after swarming as +before it. + +Thus the hive is perfectly capable of affording a second colony without +being too much weakened. The third and fourth swarm weaken it more +sensibly; but the inhabitants always remain in sufficient numbers to +preserve the course of their labours uninterrupted; and the losses are +soon repaired by the great fecundity of the queen, as she lays above an +hundred eggs a day. + +If, in some cases, the agitation of swarming is so great, that all the +bees participate in it, and leave the hive, the desertion lasts but for +a moment. The hive throws only during the finest part of the day, and it +is then that the bees are ranging through the country. Those that are +out, therefore, cannot share in the agitation; when returned to the +hive, they quietly resume their labours; and their number is not small, +for, when the weather is fine, at least a third of the bees are employed +in the fields at once. + +Even in the most embarrassing case, namely, where the whole bees desert +the hive, it does not follow, that all those endeavouring to depart +become members of the new colony. When this agitation or delirium seizes +them, the whole rush forward and accumulate towards the entrance of the +hive, and are heated in such a manner that they perspire copiously. +Those near the bottom, and supporting the weight of all the rest, seem +drenched in perspiration; their wings grow moist; they are incapable of +flight; and even when able to escape, they advance no farther than the +board of the hive, and soon return. + +Those that have lately left their cells remain behind the swarm, still +feeble, they could not support themselves in flight. Here then are also +many recruits to people what we should have thought a deserted +habitation. + + + + +LETTER X. + +_THE SAME SUBJECT CONTINUED._ + + +To preserve greater regularity in continuing the history of swarms, I +think it proper to recapitulate in a few words the principal points of +the preceding letter, and to expatiate on each, concerning the result of +new experiments, respecting which I have still been silent. + +In the first place. _If at the return of spring, we examine a hive well +peopled, and governed by a fertile queen, we shall see her lay a +prodigious number of male eggs in the course of May, and the workers +will chuse that moment for constructing several royal cells of the kind +described by M. de Reaumur._ Such is the result of several long +continued observations, among which there has not been the slightest +variation, and I cannot hesitate in announcing it as demonstrated. +However, I should here add the necessary explanation. It is necessary +that the queen, before commencing her _great_ laying of the eggs of +males, be eleven months old; when young she lays only those of workers. +A queen, hatched in spring, will perhaps lay fifty or sixty eggs of +drones in whole, but before beginning her great laying of them, which +should be two thousand in a month, she must have completed her eleventh +month in age. In the course of our experiments, which more or less +disturbed the natural state of things, it often happened that the queen +did not attain this age until October, and immediately began laying male +eggs. The workers, as if induced by some emanation from the eggs, also +adopted this time for building the royal cells. No swarm resulted +thence, it is true, because in autumn all the necessary circumstances +are absolutely wanting, but it is not less evident, that there is a +secret relation between the production of the eggs of males, and the +construction of royal cells. + +This laying commonly continues thirty days. The bees on the twentieth or +twenty-first lay the foundation of several royal cells. Sometimes they +build sixteen or twenty; we have even had twenty-seven. When the cells +are three or four lines high, the queen lays those eggs from which her +own species will come, but not the whole in one day. That the hive may +throw several swarms, it is essential that the young females conducting +them be not all produced at the same time. One may affirm, that the +queen anticipates the fact, for she takes care to allow at least the +interval of a day between every egg deposited in the cells. It is proved +by the bees knowing to close the cells the moment the worms are ready to +metamorphose to nymphs. Now, as they close all the royal cells at +different periods, it is evident the included worms are not all of an +equal age. + +The queen's belly is very turgid before she begins laying the eggs of +drones; but it sensibly decreases as she advances, and when terminated +is very small. Thus she finds herself in a condition to undertake a +journey which circumstances may prolong; thus this condition was +necessary; and as every thing is harmonious in the laws of nature, the +origin of the males corresponds with that of the females, which they are +to fecundate. + +Secondly. _When the larvæ hatched from the eggs laid by the queen, in +the royal cells, are ready to transform to nymphs, this queen leaves the +swarm conducting a swarm along with her; and the first swarm that +proceeds from the hive is uniformly conducted by the old queen._{M} I +think I can divine the reason of it. + +That there may never be a plurality of females in a hive, nature has +inspired queens with a natural horror against each other; they never +meet without endeavouring to fight, and to accomplish their mutual +destruction. Thus, the chance of combat is equal between them, and +fortune will decide to which the empire shall pertain. But if one +combatant is older than the rest, she is stronger, and the advantage +will be with her. She will destroy her rivals successively as produced. +Thus, if the old queen did not leave the hive, when the young ones +undergo their last metamorphosis, it could produce no more swarms, and +the species would perish. Therefore, to preserve the species, it is +necessary that the old queen conduct the first swarm. But what is the +secret means employed by nature to induce her departure? I am ignorant +of it. + +In this country it is very rare, though not without example, for the +swarm, led forth by the old queen, in three weeks to produce a new +colony, which is also conducted by the same old queen; and that may +happen thus. Nature has not willed that the queen shall quit the first +hive before her production of male eggs is finished. It is necessary for +her to be freed of them, that she may become lighter. Besides, if her +first occupation, on entering a new dwelling, was laying more male eggs, +still she might perish either from age or accident before depositing +those of workers. The bees in that case would have no means of replacing +her, and the colony would go to ruin. + +All these things have been with infinite wisdom foreseen. The first +operation of the bees of a swarm is to construct the cells of workers. +They labour at them with great ardour, and as the ovaries of the queen +have been disposed with admirable foresight, the first eggs she has to +lay in her new abode are those of workers. Commonly her laying +continues ten or eleven days; and at this time portions of comb +containing large cells are fabricated. It may be affirmed, that the bees +know their queen will also lay the eggs of drones; she actually does +begin to deposit some, though in much smaller number than at first; +enough however to encourage the bees to construct royal cells. Now, if +in these circumstances the weather is favourable, it is not impossible +that a second colony may be formed, and that the queen may depart at the +head of it within three weeks of conducting the first swarm. But I +repeat, the fact is rare in our climate. Let me now return to the hives +from which the queen has led the first colony. + +Thirdly. _After the old queen has conducted the first swarm from the +hive, the remaining bees take particular care of the royal cells, and +prevent the young queens successively hatched from leaving them, unless +at an interval of several days between each._ + +In the preceding letter, I have given you the detail and proof of this +fact, and I shall here add some reflexions. During the period of +swarming, the conduct or instinct of bees seems to receive a particular +modification. At all other times, when they have lost their queen, they +appropriate workers worms to replace her; they prolong and enlarge the +cells of these worms; they supply them with aliment more abundantly, and +of a more pungent taste; and by this alteration, the worms that would +have changed to common bees are transformed to queens. We have seen +twenty-seven cells of this kind constructed at once; but when finished +the bees no longer endeavour to preserve the young females from the +attacks of their enemies. One may perhaps leave her cell, and attack all +the other royal cells successively, which she will tear open to destroy +her rivals, without the workers taking any part in their defence. Should +several queens be hatched at once, they will pursue each other, and +fight until the throne remain with her that is victorious. Far from +opposing such duels, the other bees rather seem to excite the +combatants. + +Things are quite reversed during the period of swarming. The royal cells +then constructed are of a different figure from the former. They +resemble stalactites, and in the beginning are like the cup of an acorn. +The bees assiduously guard the cells when the young queens are ready for +their last metamorphosis. At length the female hatched from the first +egg laid by the old queen leaves her cell; the workers at first treat +her with indifference. But she, immediately yielding to the instinct +which urges her to destroy her rivals, seeks the cells where they are +enclosed; yet no sooner does she approach than the bees bite, pull, and +drive her away, so that she is forced to remove; but the royal cells +being numerous, scarce can she find a place of rest. Incessantly +harassed with the desire of attacking the other queens, and incessantly +repelled, she becomes agitated, and hastily traverses the different +groupes of workers, to which she communicates her agitation. At this +moment numbers of bees rush towards the aperture of the hive, and, with +the young queen at their head, depart to seek another habitation. + +After the departure of the colony, the remaining workers set another +queen at liberty, and treat her with equal indifference as the first. +They drive her from the royal cells; being perpetually harrassed, she +becomes agitated; departs, and carries a new swarm along with her. In a +populous hive this scene is repeated three or four times during spring. +As the number of bees is so much reduced, that they are no longer +capable of preserving a strict watch over the royal cells, several +females then leave their confinement at once; they seek each other, +fight, and the queen at last victorious reigns peaceably over the +republic. + +The longest intervals we have observed between the departure of each +natural swarm have been from seven to nine days. This is the time that +usually elapses after the first colony is led out by the old queen, +until the next swarm is conducted by the first young queen set at +liberty. The interval between the second and third is still shorter; and +the fourth sometimes departs the day after the third. In hives left to +themselves, fifteen or eighteen days are usually sufficient for the +throwing of the four swarms, if the weather continues favourable, as I +shall explain. + +A swarm is never seen except in a fine day, or, to speak more correctly, +at a time of the day when the sun shines, and the air is calm. Sometimes +we have observed all the precursors of swarming, disorder and agitation, +but a cloud passed before the sun, and tranquillity was restored; the +bees thought no more of swarming. An hour afterwards, the sun having +again appeared, the tumult was renewed; it rapidly augmented; and the +swarm departed. + +Bees generally seem much alarmed at the prospect of bad weather. While +ranging in the fields the passage of a cloud before the hive induces +them precipitately to return. I am induced to think they are disquieted +by the sudden diminution of light. For if the sky is uniformly obscured, +and there is no alteration in clearness or in the clouds dispelling, +they proceed to the fields for their ordinary collections, and the first +drops of a soft rain does not make them return with much precipitation. + +I am persuaded that the necessity of a fine day for swarming is one +reason that has induced nature to admit of bees protracting the +captivity of their young queens in the royal cells. I will not deny that +they sometimes seem to use this right in an arbitrary manner. However +the confinement of the queens is always longer when bad weather lasts +several days together. Here the final object cannot be mistaken. If the +young females were at liberty to leave their cradles during these bad +days, there would be a plurality of queens in the hive, consequently +combats; and victims would fall. Bad weather might continue so long, +that all the queens might at once have undergone their last +metamorphosis, or attained their liberty. One victorious over the whole +would enjoy the throne, and the hive, which should naturally produce +several swarms, could give only one. Thus the multiplication of the +species would have been left to the chance of rain, or fine weather, +instead of which it is rendered independent of either, by the wise +dispositions of nature. By allowing only a single female to escape at +once, the formation of swarms is secured. This explanation appears so +simple, that it is superfluous to insist farther on it. + +But I should mention another important circumstance resulting from the +captivity of queens; which is, that they are in a condition to fly, when +the bees have given them liberty, and by this means are capable of +profiting by the first moment of sunshine to depart at the head of a +colony. + +You well know, Sir, that all drones and workers are not in a condition +to fly for a day or two after leaving their cells. Then they are of a +whitish colour, weak, and their organs infirm. At least, twenty-four or +thirty hours must elapse before the acquisition of perfect strength, and +the development of all their faculties. It would be the same with the +females was not their confinement protracted after the period of +transformation; but we see them appear, strong, full grown, brown, and +in a better condition for flying than at any other period. I have +elsewhere observed, that constraint is used to retain the queens in +captivity. The bees solder the covering to the sides of the cell by a +cordon of wax. As I have also explained how they are fed, it need not be +repeated here. + +It is likewise a very remarkable fact, that queens are set at liberty +earlier or later according to their age. Immediately when the royal +cells were sealed, we marked them all with numbers, and we chose this +period because it indicated the age of the queens exactly. The oldest +was first liberated, then the one immediately younger, and so on with +the rest. None of the younger queens were set at liberty before the +older ones. + +I have a thousand times asked myself how the bees could so accurately +distinguish the age of their captives. Undoubtedly I should do better to +answer this question by a simple avowal of my ignorance. At the same +time, I must be permitted to state a conjecture. You will admit, that I +have not, as some authors, abused the right of giving myself up to +hypothesis; may not the humming or sound emitted by the young queens in +their cells, be one of the methods employed by nature to instruct the +bees in the age of their queens? It is certain that the female, whose +cell is first sealed, is also the first to emit this sound. That in the +next emits it sooner than the rest, and so on with those immediately +subsequent. As their captivity may continue six days, it is possible +that the bees in this space of time may forget which has emitted it +first; but it is also possible, that the queens diversify the sounds, +encreasing the loudness as they become older, and that the bees can +distinguish these variations. We have even ourselves been able to +distinguish differences in the sound, either with relation to the +succession of notes, or their intensity; and probably there are +gradations still more imperceptible that escape our organs, but may be +sensible to those of the workers. + +What gives weight to this conjecture is, that the queens brought up by +M. Schirach's method, are perfectly mute; neither do the workers form +any guard around their cells, nor do they retain them in captivity a +moment beyond the period of transformation, and, when they have +undergone it, they are allowed to combat until one has become +victorious over all the rest. Why? Because the object is only to replace +the last queen. Now, provided that among the worms reared as queens, +only one succeeds, the fate of the others is uninteresting to the bees, +whereas, during the period of swarming, it is necessary to preserve a +succession of queens, for conducting the different colonies; and to +ensure the safety of the queens, it is necessary to avert the +consequences of the mutual horror by which they are animated against +each other. Behold the evident cause of all the precautions that bees, +instructed by nature, take during the period of swarming; behold an +explanation of the captivity of females; and that the duration of their +captivity might be ascertained by the age of the young queens, it was +requisite for them to have some method of communicating to the workers +when they should be liberated. This method consists in the sound +emitted, and the variation they are able to give it. + +In spite of all my researches, I have never been able to discover the +situation of the organ which produces the sound. But I have instituted a +new course of experiments on the subject, which are still unfinished. + +Another problem still remains for solution. Why are the queens reared, +according to M. Schirach's method, mute, whilst those bred in the time +of swarming have the faculty of emitting a certain sound? What is the +physical cause of this difference? At first I thought it might be +ascribed to the period of life, when the worms that are to become queens +receive the royal food. While hives swarm, the royal worms receive the +food adapted for queens, from the moment of leaving the egg; those on +the contrary, destined for queens, according to M. Schirach's method, +receive it only the second or third day of their existence. It appears +to me that this circumstance may have an influence on the different +parts of organisation, and particularly on the organ of voice. +Experiment has not confirmed this conjecture. I constructed glass cells +in perfect imitation of royal cells, that the metamorphosis of the worms +into nymphs, and of the nymphs to queens, might be visible. These +experiments are related in a preceding letter. Into one of these +artificial cells we introduced the nymph of a worm, reared according to +M. Schirach's method, twenty-four hours before it could naturally +undergo its last metamorphosis; and we replaced the glass cell in the +hive, that the nymph might have the necessary degree of heat. Next day, +we had the satisfaction of seeing it divest itself of the spoil, and +assume its ultimate figure. This queen was prevented from escaping from +her prison; but we had contrived an aperture for her thrusting out her +trunk, and that the bees might feed her. I expected that she would have +been completely mute; but it was otherwise; for she emitted sounds +similar to those already described, therefore my conjecture was +erroneous. + +I next conceived that the queen being restrained in her motions, and in +her desire for liberty, was induced to emit certain sounds. All queens, +in this new point of view, are equally capable of emitting the sound, +but to induce them to it, they must be in a confined situation. In the +natural state, the queens that come from workers are not a single +instant in restraint; and, if they do not emit the sound, it is because +nothing impels them to it. On the other hand, those produced at the time +of swarming are induced to do so by the captivity in which they are +kept. For my own part, I give little weight to this conjecture; and +though I state it here, it is less with a view to claim merit than to +put others on a plan of discovering something more probable. + +I do not ascribe to myself the credit of having discovered the humming +of the queen bee. Old authors speak of it. M. de Reaumur cites a Latin +work published 1671, _Monarchia Femina_, by Charles Butler. He gives a +very brief abstract of this naturalist's observations, who we easily see +has exaggerated or rather disguised the truth, by mixing it with the +most absurd fancies; but it is not the less evident that Butler has +heard this peculiar humming of queens, and that he did not confound it +with the confused humming sometimes heard in hives. + +Fourthly. _The young queens conducting swarms from their native hive are +still in a virgin state._ The day after, being settled in their new +abode, they generally depart in quest of the males; and this is usually +the fifth day of their existence as queens; for two or three pass in +captivity, one in their native hive, and a fifth in their new dwelling. +Those queens that come from the worm of a worker, also pass five days in +the hive before going in quest of males. So long as in a state of +virginity, both are treated with indifference by the bees; but after +returning with the external marks of fecundation, they are received by +their subjects with the most distinguished respect. However, forty-six +hours elapse after fecundation before they begin to lay. The old queen, +which leads the first swarm in spring, requires no farther commerce with +the males, for preservation of her fecundity. A single copulation is +sufficient to impregnate all the eggs she will lay for at least two +years. + + _PREGNY, 8. September 1791._ + + +FOOTNOTES: + +{M} Schirach seems to have been aware of this fact.--T. + + + + +LETTER XI. + +_THE SAME SUBJECT CONTINUED._ + + +I have collected my chief observations on swarms in the two preceding +letters; those most frequently repeated, and of which the uniformity of +result leads me to apprehend no error. I have deduced what seem the most +direct consequences; and in all the theoretical part, I have sedulously +avoided going beyond facts. What is yet to be mentioned is more +hypothetical, but it engrosses several curious experiments. + +It has been demonstrated, that the principal motive of the young females +departing when hives swarm, is their insuperable antipathy to each +other. I have repeatedly observed that they cannot gratify their +aversion, because the workers with the utmost care prevent them from +attacking the royal cells. This perpetual opposition at length creates a +visible inquietude, and excites a degree of agitation that induces them +to depart. All the young queens are successively treated alike in hives +that are to swarm. But the conduct of the bees towards the old queen, +destined to conduct the first swarm, is very different. Always +accustomed to respect fertile queens, they do not forget what they owe +to her; they allow her the most uncontrouled liberty. She is permitted +to approach the royal cells; and if she even attempts to destroy them, +no opposition is presented by the bees. Thus her inclinations are not +obstructed, and we cannot ascribe her flight, as that of the young +queens, to the opposition she suffers. Therefore, I candidly confess +myself ignorant of the motives of her departure. + +Yet, on more mature reflection, it does not appear to me that this fact +affords so strong an objection against the general rule as I had at +first conceived. It is certain at least, that the old queens, as well as +the young ones, have the greatest aversion to the individuals of their +own sex. This has been proved by the numerous royal cells destroyed. You +will remember, Sir, that in my first observations on the departure of +old queens, seven royal cells opened at one side were destroyed by the +queen. If rain continues several days, the whole are destroyed; in this +case, there is no swarm, which too often happens in our climate, where +spring is generally rainy. Queens never attack cells containing an egg +or a very young worm; but only when the worm is ready for transforming +to a nymph, or when it has undergone its last metamorphosis. + +The presence of royal cells with nymphs or worms near their change, also +inspires old queens with the utmost horror or aversion; but here it +would be necessary to explain why the queen does not always destroy them +though it is in her power. On this point, I am limited to conjectures. +Perhaps the great number of royal cells in a hive at once, and the +labour of opening the whole, creates insuperable alarm in the old queen. +She commences indeed with attacking her rivals; but, incapable of +immediate success, her inquietude during this work becomes a terrible +agitation. If the weather continues favourable, while she remains in +this condition, she is naturally disposed to depart. + +It may easily be understood, that the workers accustomed to respect +their queen, whose presence is a real necessity to them, crowd after +her; and the formation of the first swarm creates no difficulty in this +respect. But you will undoubtedly ask, Sir, What motive can induce the +workers to follow their queen from the hive, while they treat the young +queens very ill, and, even in their most amicable moments, testify +perfect indifference towards them. Probably it is to escape the heat to +which the hive is then exposed. The extreme agitation of the females +leads them to traverse the combs in all directions. They pass through +groupes of bees, injure and derange them; they communicate a kind of +delirium, and these tumultuous motions raise the temperature to an +insupportable degree. We have frequently proved it by the thermometer. +In a populous hive it commonly stands between 92° and 97°, in a fine day +of spring; but during the tumult which precedes swarming, it rises above +104°. And this is heat intolerable to bees. When exposed to it, they +rush impetuously towards the outlets of the hive and depart. In general +they cannot endure the sudden augmentation of heat, and in that case +quit their dwelling; neither do those returning from the fields enter +when the temperature is extraordinary. + +I am certain, from direct experiments, that the impetuous courses of the +queen over the combs, actually throws the workers into agitation; and I +was able to ascertain it in the following manner. I wished to avoid a +complication of causes. It was particularly important to learn, whether +the queen would impart her agitation but not at the time of swarming. +Therefore I took two females still virgins, but capable of fecundation +for above five days, and put one in a glass hive sufficiently populous; +the other I put into a different hive of the same kind. Then I shut the +hives in such a way that there was no possibility of their escape. The +air had free circulation. I then prepared to observe the hives every +moment that the fineness of the weather would invite both males and +females to go abroad, for the purpose of fecundation. Next morning, the +weather being gloomy, no male left the hive, and the bees were tranquil; +but towards eleven of the following day, the sun shining bright, both +queens began to run about seeking an exit from every part of their +dwelling; and from their inability to find one, traversed the combs with +the most evident symptoms of disquiet and agitation. The bees soon +participated of the same disorder; they crowded towards that part of the +hive where the openings were placed; unable to escape they ascended with +equal rapidity, and ran heedlessly over the cells until four in the +afternoon. It is nearly about this time that the sun declining in the +horizon recalls the males; queens requiring fecundation never remain +later abroad. The two females became calmer, and tranquillity was in a +short time restored. This was repeated several subsequent days with +perfect similarity; and I am now convinced that there is nothing +singular in the agitation of bees while swarming, but that they are +always in a tumultuous state when the queen herself is in agitation. + +I have but one fact more to mention. It has already been observed, that +on losing the female, bees give the larvæ of simple workers the royal +treatment, and, according to M. Schirach, in five or six days they +repair the loss of their queen. In this case there are no swarms. All +the females leave their cells almost at the same moment, and after a +bloody combat the throne remains with the most fortunate. + +I can very well comprehend that the object of nature is to replace the +lost queen; but as bees are at liberty to choose either the eggs or +worms of workers, during the first three days of existence; to supply +her place, why do they give the royal treatment to worms, all of nearly +an equal age, and which must undergo their last metamorphosis almost at +the same time? Since they are enabled to retain the young females in +their cells, why do they allow all the queens, reared according to +Schirach's method, to escape at once. By prolonging their captivity +more or less, they would fulfil two most important objects at once, in +repairing the loss of their females and preserving a succession of +queens to conduct several swarms. + +At first it was my opinion, that this difference of conduct proceeded +from the difference of circumstances in which they found themselves +situated. They are induced to make all their dispositions relative to +swarming only when in great numbers, and when they have a queen occupied +with her principal laying of male eggs; whereas, having lost their +female, the eggs of drones are no longer in the combs to influence their +instinct. They are in a certain degree restless and discouraged. + +Therefore, after removing the queen from a hive, I thought of rendering +all the other circumstances as similar as possible to the situation of +bees preparing to swarm. By introducing a great many workers, I +encreased the population to excess, and supplied them with combs of +male brood in every stage. Their first occupation was to construct royal +cells after Schirach's method, and to rear common worms with royal food. +They also began some stalactite cells, as if the presence of the male +brood had inspired them to it; but this they discontinued, as there was +no queen to deposit her eggs. Finally, I gave them several close royal +cells, taken indifferently from hives preparing to swarm. However, all +these precautions were fruitless; the bees were occupied only with +replacing their lost queen; they neglected the royal cells entrusted to +their care; the included queens came out at the ordinary time, without +being detained prisoners a moment; they engaged in several combats, and +there were no swarms. + +Recurring to subtleties, we may perhaps suggest a cause for this +apparent contradiction. But the more we admire the wise dispositions of +the author of nature, in the laws he has prescribed to the industry of +animals, the greater reserve is necessary in admitting any theory +adverse to this beautiful system, and the more must we distrust that +facility of imagination from which we think by embellishment to +elucidate facts. + +In general, Naturalists who have long observed animals, and those in +particular who have chose insects for their favourite study, have too +readily ascribed to them our sentiments, our passions, and even our +intentions and designs. Incited to admiration, and disgusted perhaps by +the contempt with which insects are treated, they have conceived +themselves obliged to justify the consumption of time bestowed on this +pursuit, and they have painted different traits of the industry of these +minute animals, with the colours inspired by an exalted imagination. Nor +is even the celebrated Reaumur to be acquitted of such a charge. He +frequently ascribes combined intentions to bees; love, anticipation, and +other faculties of too elevated a kind. I think I can perceive that +although he formed very just ideas of their operations, he would be +well pleased that his reader should admit they were sensible of their +own interests. He is a painter who by a happy interest flatters the +original, whose features he depicts. On the other hand, Buffon unjustly +considers bees as mere automatons. It was reserved for you, Sir, to +establish the theory of animal industry on the most philosophical +principles, and to demonstrate that those actions that have a moral +appearance depend on an association of ideas _simply sensible_. It is +not my object here to penetrate those depths, or to insist on the +details. + +But, on the whole, facts relative to the formation of swarms perhaps +present more subjects of admiration than any other part of the history +of bees. I think it proper to state, in a few words, the simplicity of +the methods by which the wisdom of nature guides their instinct. It +cannot allow them the slightest portion of understanding; it leaves them +no precautions to be taken, no combination to be followed, no foresight +to exercise, no knowledge to acquire. But having adapted their +sensorium to the different operations with which they are charged, it is +the impulse of pleasure which leads them on. She has therefore +pre-ordained all that is relative to the succession of their different +labours; and to each operation she has united an agreeable sensation. +Thus, when bees construct cells, watch over their larvæ, and collect +provisions, we must not seek for method, affection, or foresight. The +only inducement must be sought for in some pleasing sensation attached +to each of these operations. I address a philosopher; and as these are +his own opinions applied to new facts, I believe my language will be +easily understood. But I request my readers to peruse and to reflect on +that part of your works which treats of the industry of animals. Let me +add but another sentence. The inducement of pleasure is not the sole +agent; there is another principle, the prodigious influence of which, at +least with regard to bees, has hitherto been unknown, that is the +sentiment of aversion which all females continually feel against each +other, a sentiment whose existence is so fully demonstrated by my +experiments, and which explains many important facts in the theory of +swarms. + + _PREGNY, 10. September 1791._ + + + + +LETTER XII. + +_ADDITIONAL OBSERVATIONS ON QUEENS THAT LAY ONLY THE EGGS OF DRONES, AND +ON THOSE DEPRIVED OF THE ANTENNÆ._ + + +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. + +Another very remarkable circumstance, which this experiment gave me +occasion to observe, consists in the good reception given by the bees to +the stranger queen, while they still preserved the first. Having so +often seen the symptoms of discontent that a plurality of queens +occasions, after having witnessed the clusters formed around these +supernumerary queens to confine them, I could not expect they would pay +the same homage to a second mutilated one they still testified towards +the first. Is it because after losing the antennæ, these queens have no +more any characteristic which distinguishes the one from the other? + +I was the more inclined to admit this conjecture from the bad reception +of a third fertile queen preserving her antennæ, which was introduced +into the same hive. The bees seized, bit her, and confined her so +closely, that she could hardly breath or move. Therefore, if they treat +two females deprived of antennæ in the same hive equally well, it is +probably because they experience the same sensation from these two +females, and want the means of longer distinguishing them from each +other. + +From all this, I conclude, that the antennæ are not a frivolous ornament +to insects, but, according to all appearance, are the organs of touch +or smell. Yet I cannot affirm which of these senses reside in them. It +is not impossible that they are organised in such a manner as to fulfil +both functions at once. + +As in the course of this experiment both mutilated females constantly +endeavoured to escape from the hive, I wished to see what they would do +if set at liberty, and whether the bees would accompany them in their +flight. Therefore I removed the first and third queen from the hive, +leaving the fertile mutilated one, and enlarged the entrance. + +The queen left her habitation the same day. At first she tried to fly, +but, her belly being full of eggs, she fell down and never attempted it +again. No workers accompanied her. Why, after rendering the queen so +much attention while she lived among them, did they abandon her now on +her departure? You know, Sir, that queens governing a weak swarm are +sometimes discouraged, and fly away, carrying all their little colony +along with them. In like manner sterile queens, and those whose dwelling +is ravaged by weevils, depart; and are followed by all their bees. Why +therefore in this experiment did the workers allow their mutilated queen +to depart alone? All that I can hazard on this question is a conjecture. +It appears that bees are induced to quit the hives from the increased +heat which occasions the agitation of their queen, and the tumultuous +motion which she communicates to them. A mutilated queen, +notwithstanding her delirium, does not agitate the workers, because she +seeks the uninhabited parts of the hive, and the glass panes of it: she +hurries over clusters of bees, but the shock resembles that of any other +body, and produces only a local and instantaneous motion. The agitation +arising from it, is not communicated from one place to another, like +that produced by a queen, which in the natural state wishes to abandon +her hive and lead out a swarm; there is no increased heat, consequently +nothing that renders the hive insupportable to her. + +This conjecture, which affords a tolerable explanation why bees persist +in remaining in the hive, though the mutilated queen has left it, is no +reason for the motive inducing the queen herself to depart. Her instinct +is altered; that is the whole that I can perceive. I can discern nothing +more. It is very fortunate for the hive, that this queen departs, for +the bees incessantly attend her; nor do they ever think of procuring +another while she remains; and if she was long of leaving them, it would +be impossible to replace her; for the workers worms would exceed the +term at which they are convertible into royal worms, and the hive would +perish. Observe, that the eggs dropped by the mutilated queen can never +serve for replacing her, for, not being deposited in cells, they dry and +produce nothing. + +I have yet to say a few words on females laying male eggs only. M. +Schirach supposes that one branch of their double ovary suffers some +alteration. He seems to think that one of these branches contains the +eggs of males, while the other has none but common eggs, and as he +ascribes the inability of certain queens to lay the latter to some +disease, his conjecture becomes very plausible. In fact, if the eggs of +males and workers are indiscriminately mixed in both branches of the +ovary, it appears at first sight that whatever cause acts on that organ, +it should equally affect both species of eggs. If on the contrary, one +branch is occupied by the eggs of drones only, and the other contains +none but common eggs, we may conceive how disease affects the one, while +the other remains untouched. Though this conjecture is probable, it is +confuted by observation. We lately dissected queens, which laid none but +male eggs, and found both branches of the ovary equally well expanded, +and equally sound, if I may use the expression. The only difference +that struck us was that in these two branches, the eggs were apparently +not so close together as in the ovaries of queens laying both kinds of +eggs. + + _PREGNY, 12. September 1791._ + + + + +LETTER XIII. + +_ECONOMICAL CONSIDERATIONS ON BEES._ + + +In this letter I shall treat of the advantages that may be derived from +the new invented hives, called _book_ or _leaf_ hives, in promoting the +_economical knowledge_ of bees. + +It is needless to relate the different methods hitherto employed in +forcing bees to yield up a portion of their honey and wax; all resemble +each other in being cruel and ill understood. + +It is evident, when bees are cultivated for the purpose of sharing the +produce of their labours, we must endeavour to multiply them as much as +the nature of the country admits; and consequently to regard their lives +at the time we plunder them. Therefore it is an absurd custom to +sacrifice whole hives to get at the riches they contain. The inhabitants +of this country, who follow no other method, annually lose immense +numbers of hives; and spring, being generally unfavourable to swarms, +the loss is irreparable. I well know that at first they will not adopt +any other method; they are too much attached to prejudices and old +customs. But naturalists and intelligent cultivators of bees will be +sensible of the utility of the method I propose; and if they apply it to +use I hope their example will extend and perfect the culture of bees. + +It is not more difficult to lodge a natural swarm in a leaf hive than in +any other of a different shape. But there is one precaution essential +to success, which I should not omit. Though the bees are indifferent as +to the position of their combs, and as to their greater or lesser size, +they are obliged to construct them perpendicular to the horizon, and +parallel to each other. Therefore, if left entirely to themselves, when +establishing a colony in one of those new hives, they would frequently +construct several small combs parallel indeed, but perpendicular to the +plane of the frames or leaves, and by this disposition prevent the +advantages which I think to derive from the figure of my hives, since +they could not be opened without breaking the combs. Thus they must +previously have a guide to follow; the cultivator himself lays the +foundation of their edifices, and that by a simple method. A portion of +comb must be solidly fixed in some of the boxes composing the hive; the +bees will extend it; and, in prosecution of their work, will accurately +follow the plan already given them. Therefore on opening the hive, no +obstacle is to be removed, nor stings to be dreaded, for one of the +most singular and valuable properties attending this construction, is +its rendering the bees tractable. I appeal to you, Sir, for the truth of +what I say. In your presence I have opened all the divisions of the most +populous hives, and the tranquillity of the bees has given you great +surprise. I can desire no other evidence of my assertion. It is in the +facility of opening these hives at pleasure that all the advantages lie, +which I expect in perfecting the economical knowledge of bees. + +I conceive, when I observe bees may be rendered tractable, that it need +not be added, I do not arrogate to myself the absurd pretence of +_taming_ them, for this excites a vague idea of tricks; and I would +willingly avoid the hazard of exposing myself to any such reproach. I +ascribe their tranquillity on opening the hives, to the manner that the +sudden introduction of light affects them; then, they seem rather to +testify fear than anger. Many retire and enter the cells, and appear to +conceal themselves. What confirms my conjecture is, their being less +tractable during night or after sunset than through the day. Thus, we +must open the hives, while the sun is above the horizon, cautiously, and +without any sudden shock. The divisions must be separated slowly, and +care taken not to wound any of the bees. If they cluster too much on the +combs, they must be brushed off with a feather; and breathing on them +carefully avoided. The air we expire seems to excite their fury; it +certainly has some irritating quality, for if bellows are used, they are +rather disposed to escape than to sting. + +Respecting the advantages of leaf hives, I shall observe, they are very +convenient for forming _artificial_ swarms. In the history of natural +swarms, I have shewn how many favourable circumstances are necessary for +their success. From experience I know that they very often fail in our +climate; and even when a hive is disposed to swarm, it frequently +happens that the swarm is lost either because the moment of its +departure has not been foreseen, because it rises out of sight, or +settles on inaccessible places. Instructing the cultivators of bees how +to make artificial swarms is a real service, and the form of my hives +renders this an easy operation. But it requires farther illustration. + +Since bees, according to M. Schirach's discovery, can procure another +queen after having lost their own, provided there is workers brood in +the combs not above three days old, it results that we can at pleasure +produce queens, by removing the reigning one. Therefore, if a hive +sufficiently populous is divided in two, one half will retain the old +queen, and the other will not be long of obtaining a new one. But to +ensure success, we must choose a propitious moment, which is never +certain but in leaf hives. In these we can see whether the population is +sufficient to admit of division, if the brood is of the proper age, if +males exist or are ready to be produced for impregnating the young +queens. + +Supposing the union of all these conditions, the following is the method +to be pursued. The leaf hive may be divided through the middle without +any shock. Two empty divisions may be insinuated between the halves, +which, when exactly applied to each other, are close on the outside. The +queen must be sought in one of the halves, and marked to avoid mistake. +If she by chance remains in the division with most brood, she is to be +transferred to the other with less, that the bees may have every +possible opportunity of obtaining another female. Next, it is necessary +to connect the halves together, by a cord tied tight around them, and +care must be taken that they are set on the same board that the hive +previously occupied. The old entrance, now become useless, will be shut +up; but as each half requires a new one, it ought to be made at the +bottom of each division, on purpose that they may be as far asunder as +possible. Both entrances should not be made on the same day. The bees in +the half deprived of the queen ought to be confined twenty-four hours, +and no opening made before then except for admission of air. Without +this precaution, they would soon search for their queen, and infallibly +find her in the other division. They will then retire in great numbers +from their own division, until too few remain to perform the necessary +labours. But this will not ensue if they are confined twenty-four hours, +provided that interval is sufficient to make them forget the queen. When +all these circumstances are favourable, the bees, in the division +wanting the queen, will the same day begin to labour in procuring +another, and ten or fifteen days after the operation, their loss will be +repaired. The young female they have reared, soon issues forth to seek +impregnation, and in two days commences the laying of workers eggs. +Nothing more is wanting to the bees of this half hive, and the success +of the artificial swarm is ensured. + +It is to M. Schirach that we are indebted for this ingenious method of +forming swarms. He supposes, by producing young queens early in spring, +that early swarms might be procured, which would certainly be +advantageous in favourable circumstances. But unfortunately this is +impossible. Schirach believed that queens were impregnated of +themselves, consequently he thought that after being artificially +produced, they would lay and give birth to a numerous posterity. Now, +this is an error; the females, to become fertile, require the concourse +of the males, and if not impregnated within a few days of their origin, +their laying, as I have observed, is completely deranged. Thus, if a +swarm were artificially formed before the usual time of the males +originating, the bees would be discouraged by the sterility of the young +female. Or should they remain faithful to her, awaiting the period of +fecundation, as she could not for three or four weeks receive the +approaches of the male, she would lay eggs producing males only, and the +hive in this case would perish. Thus the natural order must not be +deranged, but we must delay the division of hives until males are about +to originate or actually exist. + +Besides, if M. Schirach did succeed in obtaining artificial swarms, +notwithstanding the great inconvenience of his hives, it was owing to +his singular address and unremitting assiduity. He had some pupils in +the art; these communicated the method of forming artificial swarms to +others, and there are people now in Saxony who traverse the country +practising this operation. Those versant in the matter can alone venture +to undertake it with common hives, whereas, every cultivator can do it +himself with the leaf hives. + +In this construction, another very great advantage will also be found. +Bees can be forced to work in wax. Here I am led to what I believe is a +new observation. While naturalists have directed our admiration to the +parallel position of the combs, they have overlooked another trait in +the industry of bees, namely, the equal distance uniformly between them. +On measuring the interval separating the combs, it will generally be +found four lines. Were they too distant, it is very evident the bees +would be much dispersed and unable to communicate their heat +reciprocally; whence the brood would not be exposed to sufficient +warmth. Were the combs too close, on the contrary, the bees could not +freely traverse the intervals, and the work of the hive would suffer. +Therefore, a certain distance always uniform is requisite, which +corresponds equally well with the service of the hive, and the care +necessary for the worms. Nature, which has taught bees so much, has +instructed them regularly to preserve this distance. At the approach of +winter, they sometimes elongate the cells which are to contain the +honey, and thus contract the intervals between the combs. But this +operation is a preparation for a season, when it is important to have +plentiful magazines, and when their activity being very much relaxed, it +is unnecessary for their communications to be so spacious and free. On +the return of spring, the bees hasten to contract these elongated cells, +that they may become fit for receiving the eggs which the queen will +lay, and thus re-establish the just distance which nature has ordained. + +This being admitted, bees may be forced to work in wax, or, which is the +same thing, to construct new combs. To accomplish the object, it is only +necessary to separate those already built so far asunder that they may +build others in the interval. Suppose an artificial swarm is lodged in a +leaf hive, composed of six divisions, each containing a comb, if the +young queen is as fertile as she ought to be, the bees will be very +active in their labours, and disposed to make great collections of wax. +To induce them towards this an empty box or division must be placed +between two others, each containing a comb. As all the boxes are of +equal dimensions, and of the necessary width for receiving a comb, the +bees having sufficient space for constructing a new one in the empty +division introduced into the hive, will not fail to build it, because +they are under the necessity of never having more than four lines +between them. Without any guide, this new comb will be parallel to the +old ones, to preserve that law which establishes an equal distance +throughout the whole. + +If the hive is strong and the weather good, three empty divisions may at +first be left between the old combs; one between the first and second, +another between the third and fourth, and the last between the fifth and +sixth. The bees will fill them in seven or eight days, and the hive then +contains nine combs. Should the temperature of the weather continue +favourable, three new leaves or divisions may be introduced; +consequently in fifteen days or three weeks, the bees will have been +forced to construct six new combs. The experiment may be extended +farther in warm climates, and where flowers perpetually blow. But in our +country, I have reason to think that the labour should not be forced +more during the first year. + +From these details, you are sensible, Sir, how preferable _leaf hives_ +are to those of any other construction, and even to those ingenious +stages described by _M. Palteau_, for the bees cannot by means of them +be forced to labour more in wax than they would do if left to +themselves; whereas, they are obliged to do it by inserting empty +divisions. Next, the combs constructed on those stages cannot be removed +without destroying considerable portions of brood, deranging the bees, +and creating real disorder in the hive. + +Mine have also this advantage, that what passes within may daily be +observed, and we may judge of the most favourable moments for depriving +the bees of part of their stores. With all the combs before us we can +distinguish those containing brood only, and what it is proper to +preserve. The scarcity or abundance of provisions is visible, and the +portion suitable may be taken away. + +I should protract this letter too much, if I gave an account of all my +observations on the time proper for inspecting hives, on the rules to be +followed in the different seasons, and the proportion to be observed in +dividing their riches with them. The subject would require a separate +work; and I may perhaps one day engage in it; but until that arrives I +shall always feel gratification in communicating to cultivators, who +wish to follow my method, directions of which long practice has +demonstrated the utility. + +Here I shall only observe, that we hazard absolute ruin of the hives, +by robbing them of too great a proportion of honey and wax. In my +opinion, the art of cultivating these animals consists in moderately +exercising the privilege of sharing their labours; but as a compensation +for this, every method must be employed which promotes the +multiplication of bees. Thus, for example, if we desire to procure a +certain quantity of honey and wax annually, it will be better to seek it +in a number of hives, managed with discretion, than to plunder a few of +a great proportion of their treasures. + +It is indubitable that the multiplication of these industrious animals +is much injured by privation of several combs, in a season unfavourable +to the collection of wax, because the time consumed in replacing them is +taken from that which should be consecrated to the care of the eggs and +worms, and by this means the brood suffers. Besides, they must always +have a sufficient provision of honey left for winter, for although less +is consumed during this season, they do consume some; because they are +not torpid, as some authors have conceived.{N} Therefore if they have +not enough, they must be supplied with it, which requires great +exactness. I admit that in determining to what extent hives may be +multiplied in a particular country, it is necessary first to know how +many the country can support, which is a problem yet unsolved. It also +depends on another, the solution of which is as little known, namely the +greatest distance that bees fly in collecting their provender. Different +authors maintain, they can fly several leagues from the hive. But by the +few observations I have been able to make, this distance seems greatly +exaggerated. It appears to me that the radius of the circle they +traverse does not exceed half a league. As they return to the hive with +the greatest precipitation whenever a cloud passes before the sun, it is +probable they do not fly far. Nature which has inspired them with such +terror for a storm, and even for rain, undoubtedly restrains them from +going so far as to be too much exposed to the injuries of the weather. I +have endeavoured to ascertain the fact more positively, by transporting +to various distances bees with the thorax painted, that they might again +be recognised. But none ever returned that I had carried for twenty-five +or thirty minutes from their dwelling, while those at a shorter distance +have found their way and returned. I do not state this experiment as +decisive. Though bees do not generally fly above half a league, it is +very possible they go much farther, when flowers are scarce in their own +vicinity. A conclusive experiment must be made in vast arid or sandy +plains, separated by a known distance from a fertile region. + +Thus, the question yet remains undecided. But without ascertaining the +number of hives that any district can maintain, I shall remark that +certain vegetable productions are much more favourable to bees than +others. More hives, for example, may be kept in a country abounding +meadows, and where black grain is cultivated, than in a district of +vineyards or corn. + + * * * * * + +Here I terminate my observations on bees. Though I have had the good +fortune to make some interesting discoveries, I am far from considering +my labour finished. Several problems concerning the history of these +animals still remain unsolved. The experiments I project may perhaps +throw some light on them; and I shall be animated with much greater +hopes of success, if you, Sir, will continue your counsels and +direction. I am, with every sentiment of gratitude and respect, + + FRANCIS HUBER. + _PREGNY, 1. October 1791._ + + +FOOTNOTES: + +{N} So far from being torpid in winter, when the thermometer in the open +air is several degrees below freezing, it stands at (86) and (88°), in +hives sufficiently populous. The bees then cluster together, and move to +preserve their heat. + +Now that I am on the subject of thermometrical observations, I may +cursorily remark, that M. Dubois of Bourg en Bresse, in a memoir +otherwise valuable, is of opinion, that the larvæ cannot be hatched +below (104). I have repeatedly made the experiment with the most +accurate thermometers, and obtained a very different result. When the +thermometer rises to (104°), the heat is so much greater than the eggs +require, that it is intolerable to the bees. M. Dubois has been +deceived, I imagine, by too suddenly introducing his thermometer into a +cluster of bees, and putting them in agitation, the mercury has rose +higher than it should naturally do. Had he delayed introducing the +thermometer, he would soon have seen it fall to between 95 and 97, which +is the usual temperature of hives in summer. In August this year, when +the thermometer in the open air stood at 94, it did not rise above 99 in +the most populous hives. The bees had little motion, and a great many +rested on the board of the hive. + + + + +APPENDIX. + + +[The following passages are chiefly engrossed in the substance of the +work, but the Translator, as has already been observed, for various +reasons, judges it expedient to transfer them to an appendix. In his +opinion these very minute details rather interrupt the connexion of the +narrative, however interesting they may be considered, and they pertain +more to researches purely anatomical. + +The Translator has likewise in some instances incorporated several long +and important notes with the text; because it appears to him that they +actually belong to the substance of the treatise. These are the only +variations from the original with respect to arrangement.] + + * * * * * + +Swammerdam has given an imperfect description of the ovary of the queen. +He observes that he has never been able to find the termination of the +oviducts in the abdomen, nor any other parts excepting those which he +has described. "Notwithstanding all my exertions, I never could discover +the site of the vulva, partly because I had not all my apparatus with me +in the country, when investigating this subject, and partly from my +apprehension of injuring other parts by pressure, which I had then +occasion to examine. However, I have clearly observed a muscular +swelling of the oviduct, where approaching the last ring of the belly; +that it then contracts and afterwards dilates in becoming membranaceous. +As I was desirous of preserving the poison bag, which is situated +exactly here, along with, the muscles aiding the motion of the sting, I +could follow the oviduct no farther. However, in another female, it +appeared that the vulva is in the last ring of the abdomen, and under +the sting. The parts expanding only while the queen lays, renders it +extremely difficult to penetrate the aperture." + +We have attempted to discover what has escaped the indefatigable +Swammerdam. But his observation that the research can be made to the +greatest advantage, at the time of laying, has paved the way to us. We +have remarked that the oviduct did not issue from the body, but that the +eggs fall into a kind of cavity, where they are retained several seconds +before being laid. + +On the sixth of August, we took a very fertile queen, and holding her +gently by the wings in a supine position, the whole belly was exposed. +She seized the extremity with her second pair of legs, and curved it as +much as possible. This seeming an unfavourable position for laying, we +forced her to stretch it out. The queen, oppressed with the necessity of +laying, could no longer retain her eggs. The lower part of the last ring +then separated so far from the upper part as to leave some of the inside +discovered. In this cavity the sting lay above in its sheath. As the +queen now made new efforts, we saw an egg fall into the cavity from the +end of the oviduct. The lips then closed for several seconds; they +opened again, and, in a much shorter time, dropped the egg from the +cavity. + +From our own observations we found that the seminal fluid of drones +coagulated on exposure to the air, and from several experiments had so +little doubt on the subject, that whenever the female returned with the +external marks of fecundation, we thought we recognised it in the +whitish substance filling the sexual organs. It did not then occur to us +to dissect the females to ascertain the fact more particularly: but this +year, whether designing to neglect nothing, or to examine the distension +of the female organs, we determined to dissect several. To our infinite +surprise, what we had supposed the residue of the prolific fluid, +actually proved the genital organs of the male, which separate from his +body during copulation, and remain in the female. + +We procured a number of queens according to Schirach's method for the +purpose of dissection, and set them at liberty that they might seek the +males. The first which did so, was seized the instant she returned, and +without dissection spontaneously exhibited what we were so impatient to +behold. Examining the under part of the belly, we saw the oval end of a +white substance which distended the sexual organs. The belly was in +constant motion, by alternate extension and contraction. Already had we +prepared to sever the rings, and by dissection to ascertain the cause of +these motions; when the queen curving her belly very much, and +endeavouring to reach its extremity with her hind legs, seized the +distending substance with her claws, and evidently made an effort to +extract it. She at last succeeded, and it fell before us. We expected a +shapeless mass of coagulated fluid; what therefore was our surprise to +find it part of the same male that had rendered this queen a mother. At +first we could not credit our eyes; but after examining it in every +position, both with the naked eye, and a powerful magnifier, we +distinctly recognised it to be that part which M. de Reaumur calls the +_lenticular_ body, or the _lentil_, in the following description.{O} + + 'Opening a drone there appears a portion formed by the assemblage of + several parts, often whiter than milk. This on investigation is + found to be principally composed of four oblong pieces. The two + largest are attached to a kind of twisted cord, fig. 4. r, called by + Swammerdam the root of the penis; and he has denominated seminal + vessels, s. s. two long bodies that we are about to consider. Other + two bodies oblong like the preceding, but shorter and not half the + diameter, he calls the _vasa deferentia_, d. d. Each communicates + with one of the seminal vessels near, g. g. where they unite to the + twisted cord, r. From the other extremity proceeds a very delicate + vessel, which, after several involutions, terminates in a body, t. a + little larger, but difficult to disengage from the surrounding + tracheæ. Swammerdam considers these two bodies, t. t. the testicles. + Thus there are two parts of considerable size, communicating with + other two still thicker and longer. These four bodies are of a + cellular texture, and full of a milky fluid, which may be squeezed + out. This long twisted cord, r, to which the largest of the seminal + vessels is connected, this cord, I say, is doubtless the channel by + which the milky fluid issues. After several plications, it + terminates in a kind of bladder or fleshy sac, i. i. In different + males this part is of various length and flatness. By calling it the + _lenticular_ body, or the lentil, it receives a name descriptive of + the figure it presents in all males whose internal parts have + acquired consistency in spirit of wine. The body, l. i. is therefore + a lentil, a little thickened, of which one half, or nearly so, of + the circumference is edged along the outline by two chesnut coloured + scaly plates, e. i. A small white cord, the real edge of the lentil, + is visible, and separates them. This lentil is a little oblong, and, + for convenience, we shall ascribe two extremities to it, the + anterior and posterior. The anterior, l, next the head, is where the + canal, r, dividing the seminal vessels is inserted, and the opposite + part; i. next the anus, the posterior. The two scaly plates, e. i. + e. i, proceed from the vicinity of this last part, whence each + enlarges to cover part of the lentil. Under the broadest part of + each plate, there is a division formed by two soft points of unequal + length; the largest of which is on the circumference of the lentil. + Besides these two scaly plates, there are two others, n. n. of the + same colour, narrower, and fully one half shorter, each of which is + situated very near the preceding, and originates close to the origin + of that it accompanies, namely, at the posterior part of the lentil. + The rest of the lentil is white and membranaceous. From behind + proceeds a tube, k. a canal also white and membranaceous, but it is + difficult to judge of its diameter, for the membranes, of which it + consists, are evidently in folds. To one side of this pipe is + attached a fleshy part, p. somewhat pallet shaped, one side is + concave, and the edges plaited; the other side is convex. In certain + places the plaits rise and project from the rest of the outline, and + form a kind of rays; the pallet appears prettily figured. Though + lying with the concave side applied to the lentil, it is not fixed + to it. Swammerdam seems to consider this pallet as the + characteristic part of the male. + + 'Though the parts we have described are the most conspicuous in the + male, they are neither those which protrude first, nor when + protruded are the most remarkable. On viewing from the opposite edge + of the lentil, forming the division of the two great scaly plates, a + sac or canal, k. proceeding from the posterior part of the lentil, + there is distinctly visible the body u, which we call the arc; where + there are five transverse hairy bands of a yellow colour, while the + rest is white. This arc seems out of the membranaceous canal because + it is covered only by a very transparent membrane. One end almost + reaches the lenticular body, and the other terminates where the + membranaceous canal joins the folded yellow membranes, m. which form + a species of sac, that is applied to the sides of the aperture, + adapted for the genital organs passing through. These reddish + membranes are those that appear first on pressure, and form this + elongated portion, at whose end is a kind of hairy mask. Finally, + with the sac formed by the reddish membranes, there are connected + two appendages, c. c. of reddish yellow, and red at the end, s. + These are what appear externally like horns.{P}' + +The lenticular substance, l. i. provided with each scaly lamina, are the +only parts of those described by M. de Reaumur, that we have found +engaged in the organs of our queens. The canal, r, by Swammerdam +denominated the root of the penis, breaks in copulation; and we have +seen its fragments at the place where it unites to the end of the +lentil, l. towards the anterior extremity; but we have found no traces +of the canal, k, formed of involuted membranes, which in the body of the +male proceeds from the posterior end of the lentil, l. i. nor of the +plaited pallet, p. adhering to this canal, called by Swammerdam the +penis from its resemblance to that of other animals, though he is not of +opinion that this point, which is not perforated, can perform the +functions of a real penis, and hold the principal part in generation. +The canal, k, therefore, and all appertaining to it, must break at i, +quite close to the posterior part of the lentil, since we found no +remains of the lenticular bodies left by the fecundating males, in the +body of our females. The canal, r, which Swammerdam calls the root of +the penis, with greater reason than he was himself aware, is not +extended in the body of the male as represented by the figure here +engraved, but this long twilled canal consists of several involutions, +from the seminal vessels whence it proceeds, into the lenticular body +where it terminates, and where it conveys the fluid. This canal +therefore can extend during copulation, and allow the lenticular +substance to protrude out of the body of the males. + +It is evident this may be the case during copulation as is seen on +opening a drone, for, by endeavouring to displace the lenticular body, +the involutions of the cord disappear, and it extends much more than +necessary for the lentil to protrude from the body; and if we attempt +to separate it farther, the canal breaks at l. close to the lentil, and +at the same place where it breaks in copulation. + +By dissection two nerves are discovered, towards the origin of the +canal, r. inserted into the seminal vessels and distribute in them, and +towards the root of the penis many ramifications undoubtedly serving for +the motion of these parts. Two small parts, perceptible near the nerves, +are two ligaments for retaining the generative organs in their proper +place, so that except the root of the penis, they cannot be drawn out +without some exertion; it and the lenticular body however can protrude, +and actually do so during copulation. A certain degree of pressure +forces all these parts from the body of the male, but they spontaneously +return, and appear reversed. + +Swammerdam, and after him M. de Reaumur have admired this mechanism; +they have thought, indeed, that the return should be occasioned by the +effect of the air inflating the parts, and they supposed that the male +organs proceeded from the body, and returned during copulation, the same +as when forced out by pressure. Following their example, we have pressed +them from the body of many males; we have a thousand times witnessed +this wonderful return, which they detail with the greatest precision; +but our males never survived the operation. We have seen, as M. de +Reaumur, a few males protrude them spontaneously, even some of the parts +inverted, but at that moment they died, and were unable to retract the +parts which a pressure, most likely accidental, had forced out. Thus it +is improbable that the male organs protrude by turning out of themselves +in copulation; and the details which follow prove incontestibly, that it +is otherwise. Had not Swammerdam been prejudiced with this opinion, he +would have seen that the lenticular body can proceed from the body in +erection without reversing itself; he could have proportioned the +tortuous canal, which he calls the root of the penis; he would have seen +that, at certain times, it can be sufficiently extended to allows the +lenticular substance to protrude; he would have discovered the real use +of the scaly plates; he would have explained that of the canal k, of the +plaited pallet q, and the movements of all these parts, more admirable +perhaps than the inversion which he was the first to observe. + +Our observations incontestibly prove copulation. The portion of the +males found engaged in the body of our queens, hitherto called the +lenticular substance, may be denominated a penis both from its position +and use. The same surface is presented by it in the queen as in the body +of the male, which is proved by the position of the laminæ, e. e. +attached to the interior of the penis, when found in the queen. It is +evident, if the supposed inversion took place, the laminæ would be found +within the posterior part of the penis; and we should see them through +its membrane, by their concave side, instead of which the convex surface +is presented when in the vulva of females, the same as in the body of +the males. But what is the use of these laminæ? From their figure, +hardness, relative position with respect to each other, and their +situation at the extremity of the penis, we cannot doubt they are real +pincers. However, to ascertain the fact, we found it necessary to see +their position, and that of the penis itself in the females. For this +purpose, we prevented some of the queens from extracting the parts left +by the impregnating males, and by dissection we discovered that the +laminæ were pincers as we had conjectured. + +The penis was situated under the sting of the queens, and pressed +against the upper region of the belly. It was supported by the posterior +end, against the extremity of the vagina, or excretory canal. There we +were sensible of the motion and use of the scaly pieces. Their +extremities were separated a little more than in the male, and pressed +between them some of the female parts below the excretory canal. The +extreme minuteness of these parts prevented us from distinguishing them +clearly, but the effort necessary to separate and remove the penis from +the female, satisfied us of the use of these laminæ. + +Inspecting a male from above, the convex side of the plates, e. e. is +presented, and the summit of the angle formed by their origin. When in +the body of the female, they are in the inverse position; what was above +in the male is now below, and the extremity of the pincers directed +upwards. This makes us suspect that in copulation the male mounts on the +back of the female, but we are far from asserting it positively. It may +be asked whether that part we call the penis, is the sole part +introduced into the female during copulation? We have carefully +investigated this, and can affirm, that it is the only one of all those +described by M. de Reaumur, which has been found in our females. But we +have discovered a new part that escaped both him and Swammerdam, which +appears from the following experiment. + +Separating the lenticular substance from the excretory canal, where it +was attached, we drew along with it a white body, adhering by one +extremity, and having the other engaged in the vagina. Towards the end +of the lentil, where the substance adhered, it appeared cylindrical, +then it swelled, and again contracted, to dilate anew in a greater +degree than at first; afterwards it contracted and terminated in a +point. A powerful magnifier was required to see all this. When pulled +from the lenticular body, the part was commonly broke, and also when +extracted by the queens from themselves. The figure and situation seemed +to authorise our considering it the penis itself, and the lenticular +body only an appendage. But the last queen we examined exhibited a +peculiarity that induced us to doubt the fact, and led us to suspect +that this body is nothing else than the seminal fluid itself, moulded +and coagulated in the vagina, and which from its viscosity adheres to +the lenticular substance, and accompanies it when separated from the +vagina. In this queen was found a little extravasated white matter, near +the opening of the vagina. This, though at first liquid, soon coagulated +in the air as the seminal fluid of drones does. In separating the +lenticular body from the vagina, we drew along with it a thread which +broke near the lentil; and seemed of too little consistence for the +penis of a male. The lenticular bodies, found in our queens, appeared +larger than in the males we dissected, and we have remarked with M. de +Reaumur, that these parts are not of equal size in every male. + + * * * * * + +_Experiment 1._--On the tenth of July, we set successively at liberty +three virgin queens four or five days old. Two flew away several times; +their absence was short and fruitless. The third profited better by her +liberty; she departed thrice; the first and second time her absence was +short; but the third lasted thirty-five minutes. She returned in a very +different state; and in such as allowed no doubt of her employment, for +she exhibited the part of a male that had rendered her a mother. We +seized her wings with one hand, and in the other received the +lenticular body, of which she had disengaged herself with her claws. The +posterior part was armed with two pincers, e. e. shelly and elastic, +which could be drawn asunder, and then resumed their original position. +Towards the anterior part of the lentil appeared the fragment of the +root of the penis; this canal had broke half a line from the lenticular +body. We allowed the queen to enter her habitation, and adapted the +entrance so that she could not leave it unknown to us. + +On the seventeenth we found no eggs in the hive; the queen was as +slender as the first day; therefore the male, with which she had +copulated, had not impregnated her eggs. She was again set at liberty; +after twice departing, she returned with evidence of a second +copulation. We then confined her, and the eggs she afterwards laid +proved that the second copulation had been more successful than the +first and that there are some males more fit for impregnating queens +than others. However, it is very rare that the first copulation is +inefficient; we have only seen two that required it twice; all the rest +were impregnated by the first. + + * * * * * + +_Experiment 2._--On the eighteenth we put at liberty a virgin queen +twenty-seven days old, she departed twice. Her second absence was +twenty-eight minutes, and she returned with the proofs of copulation. We +prevented her from entering, and put her under a glass to see how she +would disengage the male organs. This she was unable to accomplish, +having only the table and sides of the glass for support; therefore we +introduced a bit of comb; thus providing the same conveniences as are in +a hive. Fixing herself on _it_ by the first four legs, she stretched out +the two last, and extending them along her belly seemed to press it +between them. At length introducing her claws between the two parts of +the last ring, she seized the lenticular body, and dropped it on the +table. The posterior part was provided with shelly pincers, under which +and in the same direction was a grey cylindrical body. The end farthest +from the lentil was sensibly thicker than that adhering to it, and +terminated in a point. This point was double, and open like the bill of +a bird, which induces us to think the body was broken, a conjecture +supported by the following experiment. + + * * * * * + +_Experiment 3._--On the nineteenth we set at liberty a queen four days +old; she departed twice; her first absence was short; the second lasted +thirty minutes, and then she returned with the marks of fecundation. As +we wished to obtain the male organs entire, it was necessary to prevent +the queen from breaking them by extracting them with her feet; we +therefore suddenly killed her, and cut off the last rings in order to +lay the vulva open. But though deprived of animation, so much life +remained in these parts that the lenticular body was thrown out +spontaneously. Under the pincers appeared the remnant of a cylindrical +body which had broken near the origin and remained in the female. This +body was very small at the origin; it afterwards sensibly enlarged; next +contracting by degrees, it terminated in a sharp point. We found the +point engaged up to the gland in the excretory canal, and the rest in +the vulva. + + * * * * * + +_Experiment 4._--We set two virgin queens at liberty on the twentieth. +The first had been abroad on the preceding days, but the scarcity of +males prevented her from being previously fecundated. She returned with +the organs of a male. We tried to prevent her from extracting them, but +she did this so expeditiously with her feet, that we could not +accomplish it. She was then allowed to enter the hive. + +The second queen departed twice. Her first absence was short as usual; +the second lasted about half an hour, and she returned impregnated. +Having killed her as suddenly as possible, we laid open the vulva. The +lenticular body was deposited as in every queen hitherto dissected; the +pincers were situated under the excretory canal. Some parts not easily +distinguishable were pressed between the laminæ, and their office seemed +to consist in forcing the extremity of the lentil to approach the +orifice of the vagina, and apply so forcibly to it that some exertion +was necessary to separate them. We previously examined them, with a very +powerful magnifier. Then a peculiarity which had escaped us was +perceptible. In drawing out the lenticular body, there proceeded from +the vagina a minute part, v. adhering to the posterior end of the +lentil, and situated below the plates. It spontaneously retracted into +the lentil, like the horns of a snail. It appeared white, very short, +and cylindrical. Under the pincers was a little half coagulated seminal +fluid at the bottom of the vulva. Though much could be expressed, there +was none pure; it was almost liquid, but soon coagulated, and formed a +whitish inorganic mass. This observation carefully made removed all our +doubts, and demonstrated that what we had taken for the penis of males +was nothing but the seminal fluid, which had coagulated and assumed the +interior figure of the vagina. The only hard part introduced by the +male, was the short cylindrical point which retracted into the lentil, +when we separated it. Its situation and office prove that it is there we +must look for the issue of the seminal fluid, if we can hope to find an +opening, when not engaged in copulation. + +We found this new part in the first drone we dissected. By pressing the +seminal vessels, the white liquid then escaped downwards to the root of +the penis r. and into the lenticular body, l. i. which became sensibly +swoln. We prevented the fluid from returning, and by new pressure of the +lentil forced it to advance. However, none escaped, but we saw at the +posterior end of the lenticular body, and under the scaly pincers, a +small white cylindrical substance, the same in appearance as that we had +found engaged in the vagina of the queen. This part retracted on +pressure, and then returned. + +I request you, Sir, while perusing this letter, to inspect the figure of +the male sexual organs published by M. de Reaumur, and which are copied +here. The descriptions are most accurate, and present a just idea of the +situation of these parts when in the male's body. We readily conceive +how they appear when left in the female by copulation. This detail will +sufficiently indicate the situation and figure of the new part I have +discovered. + +I suspect that the males perish after losing their sexual organs. But +why does nature exact so great a sacrifice? This is a mystery which I +cannot pretend to unveil. I am unacquainted with any analogous fact in +natural history, but as there are two species of insects whose +copulation can take place only in the air, namely, ephemeræ and ants, it +would be extremely interesting to discover whether their males also +lose their sexual parts, in the same circumstances, and whether, as with +drones, enjoyment in their flight is the prelude of death. + +FINIS. + + +FOOTNOTES: + +{O} Memoires sur les Abeilles, p. 450. + +{P} Such long and minute descriptions can be very imperfectly +translated; indeed they are unintelligible without microscopical +inspections of the parts themselves.--T. + + + + +ANALYTICAL INDEX. + + + Description of a hive invented by the author page 4 + Swammerdam's opinion on the fecundation of bees 8 + Sentiments of M. de Reaumur 10 + Mr Debraw's opinion 11 + Hattorf's opinion 19 + Difficulty of discovering the mode of impregnation 22 + Experiments on the subject 23 + Suggestions by M. Bonnet 34 + The queen is impregnated by copulation, which never takes place + within the hive 41 + Experiments on artificial fecundation have not succeeded 42 + The male loses the sexual organs in copulation 43 + Regarded impregnation affects the ovaries of the queen 45 + She then lays no eggs but those producing males 47 + One copulation impregnates all the eggs the queen will lay in + two years 54 + Fecundity of a queen 63 + Common bees do not transport the queen's eggs 66 + They sometimes eat them 69 + Eggs producing males are sometimes laid in royal cells 71 + Common worms may be converted into queens 77 + Operations of the bees when this is done 78 + Fertile workers sometimes exist 89 + They lay none but the eggs of males 96 + All common bees are originally females 98 + Receiving the royal food while larvæ, expands their ovaries 105 + Mutual enmity of queens 110 + The common bees seem to promote their combats 117 + A guard is constantly at the entrance of the hive 123 + What ensues when bees lose their queen 126 + Effects of introducing a stranger queen 128 + Massacre of the males 132 + It never ensues in hives deprived of queens 135 + A plurality of queens is never tolerated 142 + The queen bee is oviparous 149 + Bees seem occasionally to repose 150 + Interval between production of the egg and the perfect state + of bees 151 + Mode of spinning the coccoon 153 + That of the queen is open at one end 154 + The size of the bees is not affected by that of the cells 167 + The old queen always conducts the first swarm 173 + But never before depositing eggs in the royal cells 177 + Singular effect of a sound emitted by perfect queens 189 + The instinct of bees is affected during the period of swarming 208 + Queens are liberated from their cells according to their age 214 + The bees probably judge of this by the sound emitted 217 + Young queens conducting swarms are virgins 221 + The conduct of bees to old queens is peculiar 224 + Retarded impregnation affects the instinct of queens 241 + Amputation of the antennæ produces singular effects 245 + Advantages of the leaf hive 253 + It renders the bees tractable 256 + They may there be forced to work in wax 264 + Uniform distance between the combs 265 + Natural heat of bees 269 + Distance to which they fly 271 + Appendix 273 + Anatomical observations on the sexual organs of bees 276 + Experiments proving the copulation of the queen 290 + +ALEX. SMELLIE, Printer. + + + + +{Transcriber's notes + +The spelling in the original is sometimes idiosyncratic. It has not +been changed, but a few obvious errors have been corrected. The +corrections are listed below. + +Inconsistent spellings include: + Lusace/Lusaçe, centre/center, choose/chuse, + organisation/organization, recognise/recognize + +Unusual spellings (which have not been changed) include: + centinels, coccoon, diaphraghm, encreased, encreasing, groupes, + harrassed, inaccessible, incontestible, indispensible, moveable, + perceptible, susceptible, uncontrouled, unintelligible + +Letter I + + "secret distinctive characterestics" changed to + "secret distinctive characteristics" + +Letter II + + "the copulalation of queens" changed to + "the copulation of queens" + +Letter IV + + "The worms had spun their silk coccons" changed to + "The worms had spun their silk coccoons" + +Letter V + + "characteristics of commo nbees" changed to + "characteristics of common bees" + +Letter VI + + "The result of this rencounter" changed to + "The result of this encounter" + + "genius such as your's" unchanged. + + "observing that the antennae" changed to + "observing that the antennæ" + + "combats and disastrou scenes" changed to + "combats and disastrous scenes" + + "M. de Reamur speaks of these executions" changed to + "M. de Reaumur speaks of these executions" + +Letter IX + + "Only the few bees that not participated" changed to + "Only the few bees that had not participated" + +Letter XI + + "these tumultous motions" changed to + "these tumultuous motions" + +Letter XII + + "one antennæ" unchanged. + + "reside in them," changed to + "reside in them." + +Appendix + + "the cirumference is edged" changed to + "the circumference is edged" + + "he could have proportioned the tortous canal" changed to + "he could have proportioned the tortuous canal" + + "pressed between the laminae" changed to + "pressed between the laminæ" +} + + + + + +End of the Project Gutenberg EBook of New observations on the natural +history of bees, by Francis Huber + +*** END OF THIS PROJECT GUTENBERG EBOOK NEW OBSERVATIONS ON BEES *** + +***** This file should be named 26457-8.txt or 26457-8.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/2/6/4/5/26457/ + +Produced by Louise Pryor, Steven Giacomelli and the Online +Distributed Proofreading Team at https://www.pgdp.net (This +file was produced from images produced by Core Historical +Literature in Agriculture (CHLA), Cornell University) + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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