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Fertile Workers

It has already been remarked, that the workers are proved by dissection

to be females, all of which, under ordinary circumstances, are barren.

Occasionally, some of them appear to be more fully developed than

common, so as to be capable of laying eggs: these eggs, like those of

Queens whose impregnation has been retarded, _always produce drones_!

Sometimes, when a colony has lost its Queen, these drone-laying workers

e exalted to her place, and treated with equal respect and affection,

by the bees. Huber ascertained that these fertile workers were generally

reared in the neighborhood of the young Queens, and he thought that they

received some particles of the peculiar food or jelly on which the

Queens are reared. (See Royal Jelly.) He did not pretend to account for

the effect of retarded impregnation; and made no experiments to

determine the facts, as to the fecundation of these fertile workers.

Since the publication of Huber's work, nearly 50 years ago, no light has

been shed upon the mysteries of drone-laying Queens and workers, until

quite recently. Dzierzon appears to have been the first to ascertain the

truth on this subject; and his discovery must certainly be ranked as

unfolding one of the most astonishing facts in all the range of

animated nature. This fact seems, at first view, so absolutely

incredible, that I should not dare to mention it, if it were not

supported by the most indubitable evidence, and if I had not, (as I have

already observed,) determined to state all important and well

ascertained facts, without seeking, by any concealments, to pander to

the prejudices of conceited, and often, very ignorant Bee-Keepers.

Dzierzon advances the opinion that impregnation is not needed in order

that the eggs of the Queen may produce drones; but, that all impregnated

eggs produce females, either workers or Queens; and all unimpregnated

ones, males or drones! He states that he found drone-laying Queens in

several of his hives, whose wings were so imperfect that they could not

fly, and that on examination, they proved to be unfecundated. Hence he

concluded that the eggs of the Queen Bee or fertile worker, had from the

previous impregnation of the egg which produced them, sufficient

vitality to produce the drone, which is a less highly organized insect,

and one inferior to the Queen or workers. It had long been known, that

the Queen deposits drone eggs in the large or drone cells, and worker

eggs in the small or worker cells, and that she makes no mistakes.

Dzierzon inferred, therefore, that there was some way in which she was

able to decide as to the sex of the egg before it was laid, and that she

must have a control over the mouth of the seminal sac, so as to be able

to extrude her eggs, allowing them to receive or not, just as she

pleased, a portion of its fertilizing contents. In this way he thought

she determined the sex, according to the size of the cells in which she

laid them. Mr. Samuel Wagner of York, Pa., has recently communicated to

me a very original and exceedingly ingenious theory of his own, which he

thinks will account for all the facts without admitting that the Queen

Bee has any special knowledge or will on the subject. He supposes that

when she deposits her eggs in the worker cells, her body is slightly

compressed by the size of the cells, and that the eggs, as they pass the

spermatheca, receive in this manner, its vivifying influence. On the

contrary, when she is egg-laying in drone cells, this compression cannot

take place, the mouth of the spermatheca is kept closed, and the eggs

are, necessarily, unfecundated. This theory may prove to be true, but at

present, it is encumbered with some difficulties and requires further

investigation, before it can be considered as fully established.

Leaving then the question whether the Queen exercises any volition in

this matter, for the present undecided, I shall state some facts which

occurred in the summer of 1852, in my own Apiary, and shall then

endeavor to relieve, as far as possible, this intricate subject from

some of the difficulties which embarrass it.

In the Autumn of 1852, my assistant found, in one of my hives, a young

Queen, the whole of whose progeny was drones. The colony had been formed

by removing part of the combs containing bees, brood and eggs from

another hive. It had only a few combs, and but a small number of bees.

They raised a new Queen in the manner which will hereafter be

particularly described. This Queen had laid a number of eggs in one of

the combs, and the young bees from some of them were already emerging

from the cells. I perceived, at the first glance, that they were drones.

As there were none but worker cells in the hive, they were reared in

them, and not having space for full development, they were dwarfed in

size, although the bees, in order to give them more room, had pieced out

the cells so as to make them larger than usual! Size excepted, they

appeared as perfect as any other drones.

I was not only struck with the singularity of finding drones reared in

worker cells, but with the equally singular fact that a young Queen, who

at first lays only the eggs of workers, should be laying drone eggs at

all; and at once conjectured that this was a case of a drone-laying,

unimpregnated Queen, as sufficient time had not elapsed for her

impregnation to be unnaturally retarded. I saw the great importance of

taking all necessary precautions to determine this point. The Queen was

removed from the hive, and carefully examined. Her wings, although they

appeared to be perfect, were so paralized that she could not fly. It

seemed probable, therefore, that she had never been able to leave the

hive for impregnation.

To settle the question beyond the possibility of doubt, I submitted this

Queen to Dr. Joseph Leidy for microscopic examination. The following is

an extract from his report: "The ovaries were filled with eggs; the

poison sac was full of fluid, and I took the whole of it into my mouth;

the poison produced a strong metallic taste, lasting for a considerable

time, and at first, it was pungent to the tip of the tongue. The

spermatheca was distended with a perfectly colorless, transparent,

viscid liquid, _without a trace of spermatozoa_."

This examination seems perfectly to sustain the theory of Dzierzon, and

to demonstrate that Queens do not need to be impregnated, in order to

lay the eggs of males.

I must confess that very considerable doubts rested on my mind, as to

the accuracy of Dzierzon's statements on this subject, and chiefly

because of his having hazarded the unfortunate conjecture that the place

of the poison bag in the worker, is occupied in the Queen, by the

spermatheca. Now this is so completely contrary to fact, that it was a

very natural inference that this acute and thoroughly honest observer,

made no microscopic dissections of the insects which he examined. I

consider myself peculiarly fortunate in having enjoyed the benefit of

the labors of a Naturalist, so celebrated as Dr. Leidy, for microscopic

dissections. The exceeding minuteness of some of the insects which he

has completely figured and described, almost passes belief.

On examining this same colony a few days later, I obtained the most

satisfactory evidence that these drone eggs were laid by the Queen which

had been removed. No fresh eggs had been deposited in the cells, and the

bees, on missing her, had commenced the construction of royal cells, to

rear if possible, another Queen, a thing which they would not have done,

if a fertile worker had been present, by which the drone eggs had been


Another very interesting fact proves that _all_ the eggs laid by this

Queen, were drone eggs. Two of the royal cells were, in a short time,

discontinued, and were found to be empty, while a third contained a

worm, which was sealed over the usual way, to undergo its changes from a

worm to a perfect Queen.

I was completely at a loss to account for this, as the bees having an

unimpregnated drone-laying Queen, ought not to have had a single female

egg from which they could rear a Queen.

At first I imagined that they might have _stolen_ it from another hive,

but when I opened this cell, it contained, instead of a queen, _a dead


I then remembered that Huber has described the same mistake on the part

of some of his bees. At the base of this cell, was an extraordinary

quantity of the peculiar jelly or paste, which is fed to the young that

are to be transformed into queens. The poor bees in their desperation,

appear to have dosed the unfortunate drone to death: as though they

expected by such liberal feeding, to produce some hopeful change in his

sexual organization!

It appears to me that these facts constitute all the links in a perfect

chain, and demonstrate beyond the possibility of doubt, that

unfecundated queens are not only capable of laying eggs, (this would be

no more remarkable than the same occurrence in a hen,) but that these

eggs are possessed of sufficient vitality to produce drones. Aristotle,

who flourished before the Christian era, had noticed that there was no

difference in appearance, between the eggs producing drones and those

producing workers; and he states that drones only are produced in hives

which have no queen; of course the eggs producing them, were laid by

fertile workers. Having now the aid of powerful microscopes, we are

still unable to detect the slightest difference in size or appearance in

the eggs, and this is precisely what we should expect if the same egg

will produce either a worker or a drone, according as it is or is not

impregnated. The theory which I propose, will, I think, perfectly

harmonize with all the observed facts on this subject.

I believe that after fecundation has been delayed for about three weeks,

the mouth of the spermatheca becomes permanently closed, so that

impregnation can no longer be effected; just as the parts of a flower,

after a certain time, wither and shut up, and the plant is incapable of

fructification. The fertile drone-laying workers, are in my opinion,

physically incapable of being impregnated. However strange it may

appear, or even improbable, that an unimpregnated egg can give birth to

a living being, or that the sex can be dependent on impregnation, we are

not at liberty to reject facts, because we cannot comprehend the reasons

of them. He who allows himself to be guilty of such folly, if he seeks

to maintain his consistency, will be plunged, sooner or later, into the

dreary gulf of atheism. Common sense, philosophy and religion alike

teach us to receive all undoubted facts in the natural and the

spiritual world, with becoming reverence; assured that however

mysterious to us, they are all most beautifully harmonious and

consistent in the sight of Him whose "understanding is infinite."

There is something analogous to these wonders in the bee, in what takes

place in the aphides or green lice which infest our rose bushes and

other plants. We have the most undoubted evidence that a fecundated

female gives birth to other females, and they in turn to others still,

all of which, without impregnation, are able to bring forth young, until

at length, after a number of generations, perfect males and females are

produced, and the series starts anew!

The unequaled facilities, furnished by my hives, have seemed to render

it peculiarly incumbent on me, to do all in my power to clear up the

difficulties in this intricate and yet highly important branch of

Apiarian knowledge. All the leading facts in the breeding of bees ought

to be as well known to the bee keeper, as the same class of facts in the

rearing of his domestic animals. A few crude and hasty notions, but half

understood and half digested, will answer only for the old fashioned bee

keeper, who deals in the brimstone matches. He who expects to conduct

bee keeping on a safe and profitable system, must learn that on this, as

on all other subjects, "knowledge is power."

The extraordinary fertility of the queen bee has already been noticed.

The process of laying has been well described by the Rev. W. Dunbar, a

Scotch Apiarian.

"When the queen is about to lay, she puts her head into a cell, and

remains in that position for a second or two, to ascertain its fitness

for the deposit which she is about to make. She then withdraws her

head, and curving her body downwards,[2] inserts the lower part of it

into the cell: in a few seconds she turns half round upon herself and

withdraws, leaving an egg behind her. When she lays a considerable

number, she does it equally on each side of the comb, those on the one

side being as exactly opposite to those on the other as the relative

position of the cells will admit. The effect of this is to produce the

utmost possible concentration and economy of heat for developing the

various changes of the brood!"

Here as at every step in the economy of the bee our minds are filled

with admiration as we witness the perfect adaptation of means to ends.

Who can blame the warmest enthusiasm of the Apiarian in view of a

sagacity which seems scarcely inferior to that of man.

"The eggs of bees," I quote from the admirable treatise of Bevan, "are

of a lengthened oval shape, with a slight curvature, and of a bluish

white color: being besmeared at the time of laying, with a glutinous

substance,[3] they adhere to the bases of the cells, and remain

unchanged in figure or situation for three or four days; they are then

hatched, the bottom of each cell presenting to view a small white worm.

On its growing so as to touch the opposite angle of the cell, it coils

itself up, to use the language of Swammerdam, like a dog when going to

sleep; and floats in a whitish transparent fluid, which is deposited in

the cells by the nursing-bees, and by which it is probably nourished; it

becomes gradually enlarged in its dimensions, till the two extremities

touch one another and form a ring. In this state it is called a larva or

worm. So nicely do the bees calculate the quantity of food which will be

required, that none remains in the cell when it is transformed to a

nymph. It is the opinion of many eminent naturalists that farina does

not constitute the sole food of the larva, but that it consists of a

mixture of farina, honey and water, partly digested in the stomachs of

the nursing-bees."

"The larva having derived its support, in the manner above described,

for four, five or six days, according to the season," (the development

being retarded in cool weather, and badly protected hives,) "continues

to increase during that period, till it occupies the whole breadth and

nearly the length of the cell. The nursing bees now seal over the cell,

with a light _brown cover_, externally more or less _convex_, (the cap

of a drone cell is more convex than that of a worker,) and thus

differing from that of a honey cell which is _paler_ and somewhat

_concave_." The cap of the brood cell appears to be made of a mixture of

bee-bread and wax; it is not air tight as it would be if made of wax

alone; but when examined with a microscope it appears to be reticulated,

or full of fine holes through which the enclosed insect can have air for

all necessary purposes. From its texture and shape it is easily thrust

off by the bee when mature, whereas, if it consisted wholly of wax, the

young bee would either perish for lack of air, or be unable to force its

way into the world! Both the material and shape of the lids which seal

up the honey cells are different, because an entirely different object

was aimed at; they are of pure wax to make them air tight and thus to

prevent the honey from souring or candying in the cells! They are

concave or hollowed inwards to give them greater strength to resist the

pressure of their contents!

To return to Bevan. "The larva is no sooner perfectly inclosed than it

begins to line the cell by spinning round itself, after the manner of

the silk worm, a whitish silky film or cocoon, by which it is encased,

as it were, in a pod. When it has undergone this change, it has usually

borne the name of _nymph_ or _pupa_. The insect has now attained its

full growth, and the large amount of nutriment which it has taken serves

as a store for developing the perfect insect."

"The _working bee nymph_ spins its cocoon in thirty-six hours. After

passing about three days in this state of preparation for a new

existence, it gradually undergoes so great a change as not to wear a

vestige of its previous form, but becomes armed with a firmer mail, and

with scales of a dark brown hue. On its belly six rings become

distinguishable, which by slipping one over another enables the bee to

shorten its body whenever it has occasion to do so.

"When it has reached the twenty-first day of its existence, counting

from the moment the egg is laid, it comes forth a perfect winged insect.

The cocoon is left behind, and forms a closely attached and exact lining

to the cell in which it was spun; by this means the breeding cells

become smaller and their partitions stronger, the oftener they change

their tenants; and may become so much diminished in size as not to admit

of the perfect development of full sized bees."

"Such are the respective stages of the working bee: those of the royal

bee are as follows: she passes three days in the egg and is five a worm;

the workers then close her cell, and she immediately begins spinning her

cocoon, which occupies her twenty four hours. On the tenth and eleventh

days and a part of the twelfth, as if exhausted by her labor, she

remains in complete repose. Then she passes four days and a part of the

fifth as a nymph. It is on the sixteenth day therefore that the perfect

state of queen is attained."

"The drone passes three days in the egg, six and a half as a worm, and

changes into a perfect insect on the twenty-fourth or twenty-fifth day

after the egg is laid."

"The _development_ of _each species_ likewise proceeds more slowly when

the colonies are weak or the air cool, and when the weather is very cold

it is entirely suspended. Dr. Hunter has observed that the eggs, worms

and nymphs all require a heat above 70 deg. of Fahrenheit for their


In the chapter on protection against extremes of _heat_ and _cold_, I

have dwelt, at some length, upon the importance of constructing the

hives in such a manner as to enable the bees to preserve, as far as

possible, a uniform temperature in their tenement. In thin hives exposed

to the sun, the heat is sometimes so great as to destroy the eggs and

the larvae, even when the combs escape from being melted; and the cold is

often so severe as to check the development of the brood, and sometimes

to kill it outright.

In such hives, when the temperature out of doors falls suddenly and

severely, the bees at once feel the unfavorable change; they are obliged

in self defence to huddle together to keep warm, and thus large portions

of the brood comb are often abandoned, and the brood either destroyed at

once by the cold, or so enfeebled that they never recover from the

shock. Let every bee keeper, in all his operations, remember that brood

comb must never be exposed to a low temperature so as to become chilled:

the disastrous effects are almost as certain, as when the eggs of a

setting hen are left, for too long a time, by the careless mother. The

brood combs are never safe when taken for any considerable time from the

bees, unless the temperature is fully up to summer heat.

"[4]The young bees break their envelope with their teeth, and assisted,

as soon as they come forth, by the older ones, proceed to cleanse

themselves from the moisture and exuviae with which they were surrounded.

Both drones and workers on emerging from the cell are, at first grey,

soft and comparatively helpless so that some time elapses before they

take wing.

"With respect to the cocoons spun by the different larvae, both workers

and drones spin _complete cocoons_, or inclose themselves on every side;

royal larvae construct only _imperfect cocoons_, open behind, and

enveloping only the head, thorax, and first ring of the abdomen; and

Huber concludes, without any hesitation, that the final cause of their

forming only incomplete cocoons is, that they may thus be exposed to the

mortal sting of the first hatched queen, whose instinct leads her

instantly to seek the destruction of those who would soon become her


"If the royal larvae spun complete cocoons, the stings of the queens

seeking to destroy their rivals might be so entangled in their meshes

that they could not be disengaged. 'Such,' says Huber, 'is the

instinctive enmity of young queens to each other, that I have seen one

of them, immediately on its emergence from the cell, rush to those of

its sisters, and tear to pieces even the imperfect larvae. Hitherto

philosophers have claimed our admiration of nature for her care in

preserving and multiplying the species. But from these facts we must now

admire her precautions in exposing certain individuals to a mortal


The cocoon of the royal larva is very much stronger and coarser than

that spun by the drone or worker, its texture considerably resembling

that of the silk worm's. The young queen does not come forth from her

cell until she is quite mature; and as its great size gives her abundant

room to exercise her wings she is capable of flying as soon as she quits

it. While still in her cell she makes the fluttering and piping noises

with which every observant bee keeper is so well acquainted.

Some Apiarians have supposed that the queen bee has the power to

regulate the development of eggs in her ovaries, so that few or many are

produced, according to the necessities of the colony. This is evidently

a mistake. Her eggs, like those of the domestic hen, are formed without

any volition of her own, and when fully developed, must be extruded. If

the weather is unfavorable, or if the colony is too feeble to maintain

sufficient heat, a smaller number of eggs are developed in her ovaries,

just as unfavorable circumstances diminish the number of eggs laid by

the hen; if the weather is very cold, egg-laying usually ceases

altogether. In the latitude of Philadelphia, I opened one of my hives on

the 5th day of February, and found an abundance of eggs and brood,

although the winter had been an unusually cold one, and the temperature

of the preceding month very low. The fall of 1852 was a warm one, and

eggs and brood were found in a hive which I examined on the 21st of

October. Powerful stocks in well protected hives contain some brood, at

least ten months in the year; in warm countries, bees probably breed,

every month in the year.

It is highly interesting to see in what way the supernumerary eggs of

the queen are disposed of. When the number of workers is too small to

take charge of all her eggs, or when there is a deficiency of bee bread

to nourish the young, (See chapter on Pollen,) or when, for any reason,

she judges it not best to deposit them in cells, she stands upon a comb,

and simply extrudes them from her oviduct, and the workers devour them

as fast as they are laid! This I have repeatedly witnessed in my

observing hives, and admired the sagacity of the queen in economizing

her necessary work after this fashion, instead of laboriously depositing

the eggs in cells where they are not wanted. What a difference between

her wise management and the stupidity of a hen obstinately persisting to

set upon addled eggs, or pieces of chalk, and often upon nothing at all.

The workers eat up also all the eggs which are dropped, or deposited out

of place by the queen; in this way, nothing goes to waste, and even a

tiny egg is turned to some account. Was there ever a better comment upon

the maxim? "Take care of the pence, and the pounds will take care of


Do the workers who appear to be so fond of a tit-bit in the shape of a

new laid egg, ever experience a struggle between their appetites and the

claims of duty, and does it cost them some self denial to refrain from

making a breakfast on a fresh laid egg? It is really very difficult for

one who has carefully watched the habits of bees, to speak of his little

favorites in any other way than as though they possessed an intelligence

almost, if not quite, akin to reason.

It is well known to every breeder of poultry, that the fertility of a

hen decreases with age, until at length, she becomes entirely barren; it

is equally certain that the fertility of the queen bee ordinarily

diminishes after she has entered upon her third year. She sometimes

ceases to lay Worker eggs, a considerable time before she dies of old

age; the contents of the spermatheca are exhausted; the eggs can no

longer be impregnated and must therefore produce drones.

The queen bee usually dies of old age, some time in her fourth year,

although instances are on record of some having survived a year longer.

It is highly important to the bee keeper who would receive the largest

returns from his bees, to be able, as in my hives, to catch the queen

and remove her, when she has passed the period of her greatest

fertility. In the sequel, full directions will be given, as to the

proper time and mode of effecting it.

Before proceeding farther in the natural history of the queen bee, I

shall describe more particularly, the other inmates of the hive.