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{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.