SOME STRANGE MARRIAGE-CUSTOMS: AND VIRGIN BIRTHS
The Courtship of the Cuttle-fish—The Sumptuous Cradle of the
Argonaut—The Love-darts of the Snail—Hermaphrodites and the Dangers
of Self-fertilization—Oysters and Beauty—Sex reduced to its Lowest
Terms—Parthenogenesis and Virgin Birth—The Story of the Hive-bee—The
Departure of the Queen—The New Queen and her Marriage-flight—The
Celebration of the Nuptials and its Surprising Sequel—The Widowed Queen
turns Executioner—The Queen as Mother—The Queen’s Daughters—Nursemaids’
Duties—Change of Work—The Drones and their Career—Food and Sex—The
Bumble-bee and its Life-story.
That the psychical emotions sway the goad of sexual instincts in the
higher animals there can be no doubt; and there can be as little
uncertainty that this stimulating and controlling factor gradually
loses force as we descend in the scale of animal life. Just where
it ceases it is impossible to say. A vague, nebulous intelligence
doubtless persists after these more subtle emotions have ceased, and
this, probably, in turn, gives place to purely instinctive behaviour.
These various phases of the sexual problem grade one into the other.
But they are all parts of a continuous sequence, beginning, apparently,
in relatively simple responses to chemical interactions of the kind
known as chemotaxis and ending with the passion which, in the human
race, may become a consuming fire, purifying and ennobling, or exactly
the reverse—according to the nature of the inflammable material. That
is to say, in the phenomena of sex one sees emotions in the making. The
begetting of children becomes the underlying goal of life, the hidden
heart and soul of animated nature.
This being so, one cannot but feel surprised at the discovery that,
in certain groups of the animal kingdom one meets with a strange
exception to this great rule. And this is furnished by the phenomenon
of parthenogenesis, wherein sexual desire has been dethroned.
Offspring result from Virgin births: parental care is non-existent.
This anomalous condition must be regarded as an offshoot of the
normal course of events traced in these pages, and not as a primitive
condition. This interpretation seems to be shown clearly enough in
that almost every case where parthenogenesis obtains, males, sooner
or later, make their appearance—periodically or sporadically. Every
stage between the normal, seasonal appearance of males and their entire
suppression can be traced, and an analysis of these cases demonstrates
unequivocally the uplifting character of the bi-sexual state, if only
by the fact that the uni-sexual condition makes no demands on the
parent, and does nothing to foster the growth of the higher emotions.
No attempt need now be made to discover the origin of parthenogenesis.
Let it be assumed, for the moment, that it is a condition derived from
hermaphroditism, wherein each individual is monœcious or bi-sexual.
In all diœcious or uni-sexual animals, that is to say, where the
individuals composing the species are either male or female, each
contains a leaven of the opposite sex, even when adult. It is still
a moot point whether, in the earlier stages of development, chance
decides whether the sex shall be male or female, or, at any rate,
whether the growing body is potentially male or female, till the die is
cast by some as yet undiscovered factor; or whether this is determined
from the very beginning of germinal life. In many of the lower animals,
as among the Mollusca and some of the insects, each individual is as
much male as female, and it is from a condition such as this that
parthenogenesis probably had its rise.
These two groups are selected here because they, more than any others
in like case, afford some extremely interesting gradations in this
strange phenomena of what is to be regarded as the degeneration of
sexual individuality, for each contains some members wherein the sexes
are separate, and in these cases sexual desire is present in varying
degrees. In some it is associated with very remarkable phenomena.
Among the Mollusca the Octopuses afford one of the most striking
illustrations of such phenomena. In these creatures one of the
sucker-bearing arms is more or less completely transformed to subserve
the ends of sexual congress. Without entering into the technical
details of the changes, it will suffice to remark that it is modified
in such a way as to allow the transference of the spermatozoa from
the body cavity wherein they are formed, to the arm near, or at, the
tip of which they are stored in a special sac or “spermatophore,” and
such modified arms are said to be “hectocotylized.” This extraordinary
modification attains its maximum development in the celebrated
Argonaut, and one or two of the more typical Octopuses. In the Argonaut
this arm does not make its appearance until sexual maturity has been
attained, when a large more or less globular swelling appears,
enclosing the third arm of the left side, coiled upon itself. Having
attained its full development the sac bursts and releases the arm. The
folds which formed the sac now bend back to form a new receptacle into
which the spermatophore is passed. But this is not all. The tip of the
newly released arm bears another sac, which sooner or later bursts,
forming a long, slender penis, and along the central tube of this the
spermatozoa pass from the spermatophore to their destination. Their
conveyance thereto forms the last and most amazing feature of this
strange history. The male, eager with pent-up desire, and glowing with
all the colours of the rainbow, gradually approaches the female of his
choice, who apparently awaits him with no little palpitation, and then,
with a sudden rush flings himself upon her, and apparently thrusts
the penis into her mantle cavity, when at once the whole arm breaks
off from his body and remains attached to her person, retaining its
vitality, strange as it may seem, for some considerable time, during
which, no doubt, the spermatozoa are slowly making their way out of
the spermatophore and along the channel prepared for their reception.
That the Cuttle-fish are polyandrous there seems to be little room for
doubt, inasmuch as no less than four such detached arms have been found
beneath the mantle of one female. With the majority of the Cuttle-fish
and Octopus tribe the arm is not detached, but when it is so, and this
occurs in all the species belonging to three different genera, a new
arm is grown.
[Illustration: Plate 41.
SOME REMARKABLE METHODS OF “COURTSHIP.”
1. The female Argonaut and her egg-casket.
2 and 3. The male Argonaut and his “hectocotylized” arm.
4. A Cuttle-fish (_Ocyhöe catenulata ♂_), showing the “hectocotylized”
arm described in the text, and the “spermatophore” at the base of the
long filament.
Face page 268.]
As a rule, among these animals the males are smaller than the females.
In the case of the Argonaut there is a yet more striking difference,
for the female possesses a very beautiful shell in which she carries
her eggs. This remarkable cradle, translucent and beautifully
sculptured, she attaches to her person by means of a pair of arms which
are expanded to form great lobes, almost but not quite completely
covering the shell. The earlier naturalists believed that this shell
served as a boat, and that the lobated arms were spread as sails! This
supposed fact naturally caught the fancy of the poets, who seized upon
it to point a moral and adorn a tale. Byron celebrated these imaginary
feats of seamanship in the familiar lines:
The tender Nautilus who steers his prow,
The sea-born sailor of his shell-canoe.
and Pope bids us:
Learn of the little Nautilus to sail,
Spread the thin oar, and catch the driving gale.
Sir Richard Owen years ago, however, dispelled these pretty fancies,
though the facts are surely as wonderful as the fables they have
replaced. They afford, too, one of the most striking secondary sexual
characters to be met with among the Mollusca; nowhere else, indeed,
among the members of this group is so strange a cradle to be met with.
But little, unfortunately, is known of the behaviour of these animals,
which are by far the most active of the Mollusca, and which also
display no small degree of intelligence. Their eyes, which are of great
size and complex structure, are undoubtedly far more effective organs
of vision than are possessed by any other Molluscs. It is possible,
therefore, that the sexes discover one another by sight; and it is
certain that something in the nature of a “Courtship” takes place.
The majority of the species, also, possess the most extraordinary
powers of changing their coloration, especially during moments of great
excitement. The magnificence of the hues which succeed one another,
like a series of variegated blushes suffusing the whole body, may be
one of the weapons in the armoury of Cuttle-fish love-making. In how
far the “courtship” of the Cuttle-fish resembles that of terrestrial
animals, however, is a matter on which at present nothing is really
known. That even the comparatively sedentary species, like the Octopus,
seize upon and hold territory is very improbable, for there is no
need of such landed estates, inasmuch as the offspring are not tended
and fed by the parents—this would indeed be a laborious task in the
case of some of the “Squids” which lay between thirty thousand and
forty thousand eggs! Having regard to the fact that the records of
the reproductive habits of the Octopus tribe date back to the time of
Aristotle, more than two thousand two hundred years ago—for he first
drew attention to the hectocotylized arm—it is curious that so little
has been gleaned during this vast space of time.
There are facts in regard to the sexual relationships of some of the
Snails that are in nowise less remarkable than those just related of
the Octopus tribe. Unlike the Octopuses, the Snails are hermaphrodite,
nevertheless sexual congress takes place as with unisexual species:
the eggs of the one being fertilized by the spermatozoa of the other.
During this process the orgasm of the sexual act appears to be brought
about by stabbing one another by means of a little dart formed of
carbonate of lime, the dart burying itself in the flesh and apparently
promoting a pleasurable, tingling sensation in the course of its
journey. Speedily, no doubt, it becomes absorbed, the material being
then available for the formation of a new dart.
This remarkable instrument, which is known as a “Love-dart,” or
_Spiculum amoris_, assumes a different form in each species in which
it occurs. In some the shaft is ridged like a bayonet, as in the case
of the Garden Snail, in others the form assumed is that of an awl.
These darts are formed within a special receptacle, or “dart-sac,” but
so far no explanation as to the origin of these remarkable structures
has even been hinted at. They do not seem to have been derived by the
modification of some pre-existing organ serving a different function,
as wings, for example, are derived from walking limbs, or as lungs are
derived from air-sacs. Their origin is as mysterious as their use:
for they are not found in all Snails, though they occur in one or two
Slugs—which are degenerate Snails. But no other Molluscs save the
Snails and one or two of their immediate allies are so armed.
The hermaphrodite conditions of these animals, as with other Mollusca
in like case, present some knotty points for consideration, and
especially in regard to the problem of sex-attraction. Where each
individual is as much male as female, which is the dominating factor in
desire, the maleness or the femaleness? Though each individual contains
both ova and sperm cells, probably these ripen at different times, to
avoid danger of self-fertilization. In this case the sex impulses are
on the same footing as in the case of animals wherein the sexes are
not thus combined. That is to say, the individual which is for the
moment only potentially male mates with another for the moment only
potentially female. But this being so, how does each discover the
condition of the other?
Many of the Snails, like _Helix nemoralis_, are gaily coloured. Are
these hues, these bands of black and yellow, the product of “sexual
selection”—the outcome of a process of selection from among the most
conspicuously coloured individuals as postulated by the Darwinian
theory of Sexual Selection? If so, then this choice must be regarded
as a periodic recurrence coinciding with the period during which the
individual is dominated by its female attributes. In due course it
becomes, for the time, a male, and may find itself rejected, owing
to a lack of intensity in its coloration, or, on the other hand, it
may vanquish a rival by its very splendour. Each, in short, would
help materially in this process of beautification. If the choice of
mating for it is this rather than a choice of mates—proceeds on these
lines, the bright coloration of the members of this species becomes
easy to understand. But does it? It is more than doubtful whether the
eyes of Snails are sufficiently good to distinguish the coloration
of their neighbours’ shells, or for the matter of that of their own,
for their eyes being carried on long mobile stalks, they should have
no difficulty in contemplating their own charms. And what of Snails
of more sober hues? It seems highly probable that here, as in so
many cases, scent is the selecting factor, and the coloration is an
“accidental” feature. That the colour of the shell plays no such part
as that just postulated may be gathered from the evidence afforded by
many marine species, whose shells, though conspicuously marked, are,
during life, completely enveloped and concealed by the all-investing,
fleshy mantle. In like manner the exquisite beauty in the form and
sculpturing of the shell which so many species exhibit, are characters
which cannot be regarded as due to sexual selection.
As touching the danger of self-fertilization to which reference has
been made. That this is real is shown by the fact that the ova and
spermatozoa are rarely ripe in one individual at the same time.
However, among the pulmonata, or air-breathing gastropods, it seems to
have been established that self-fertilization can, and does, occur.
That in some species, at any rate, where cross-fertilization, for some
reason, is impossible, the individual thus isolated can store up its
own spermatozoa to be used in fertilizing its own eggs. But the fact
that this rarely happens is testimony enough that such occurrences are
inimical to well-being.
The Lamellibranch, or bivalve Mollusca, _e.g._, Oyster, Mussel, and
Cockle, afford valuable evidence as to excrescences and extravagances
of growth which appeal to our eyes as ornamental, and therefore likely
to be due to the influence of sexual selection. And this because such
ornamentation is a very conspicuous feature among these animals. Yet,
save in a few cases, locomotion is impossible, and sight is wanting.
Light-distinguishing organs, and therefore eyes, are possessed by some,
but in no case probably are they strong enough to appreciate form.
Even if they did, such revelations of beauty would play no part in
mate selection from among the most ornamental; for these creatures are
commonly fixed throughout life in one position, often, indeed, buried
in mud or sand. Some move laboriously: a few, like the Cockles and
Pectens, swim by rapidly opening and closing the shell. The Pectens
are brilliantly coloured, not only as regards the shell, which is also
beautifully sculptured, but the foot also is of a vivid scarlet, and
the Pecten have numerous minute eyes. But the Cockles and Mussels
possess like attributes as to colour and sculpture, yet they are
blind. More to the point is the fact that these animals do not mate
after the fashion of higher animals, but the males, where the sexes
are distinct, discharge immense quantities of spermatozoa into the
water, and these find their way to the ova of the female through the
action of the inhalent currents set up by the animal for the purpose of
drawing in fresh supplies of water containing food and oxygen. There
are no “secondary sexual characters,” that is to say, that even where
the sexes are separate, and many, like the Oysters, are hermaphrodite,
they are externally indistinguishable. Nevertheless, many, as has been
already remarked, have shells of great beauty. As, for example, the
giant Tridacna and the strangely spinous valves of the “Thorny Oysters”
(_Spondylidæ_).
The fact that the Lamellibranch, or bivalve molluscs, are far less
numerous in point of species than the univalve tribes is accounted for
by the fact that in the first place they are of necessity aquatic, and
in the second their means of locomotion is extremely limited. Some few
species swim spasmodically: some crawl: many are incapable of movement
when once the motile larva settles down and the shell-bearing adult
stage is attained. Such species can extend their range only by means
of larval wanderings. Enormous numbers, millions, of young have to be
produced and set adrift each year by every adult in the community, and
yet but a few of each brood can ever attain to maturity. Life, for such
species, must be a dull, monotonous business: the only opportunity for
excitement is that which is preliminary to being eaten, and the only
purpose in life is to be eaten. But happily Oysters don’t think. They
and their kind are simply semi-conscious living things, responding
mechanically to stimuli. Any approach, then, to beauty, either of form
or coloration, or both, must be regarded as due to innate, inherent
changes in the germ-plasm affecting the parts so made conspicuous:
the only form of selection to which such “ornaments” can be subjected
is Natural Selection. If, and when, such ornaments penalize their
possessor either by their cumbrousness or their conspicuous characters,
or by increasing the difficulty of feeding or distributing offspring,
then the further development of such excrescences is checked by the
death of all individuals which have passed the bounds of endurance in
this respect.
Sex, and all that appertains thereto, in short, is in these creatures
reduced to its lowest terms. There are not wanting, to-day, both men
and women, who affect to believe that all would be well for the human
race could a similar slowing-down, or strangulation, of the sexual
instincts be brought about. Such blind leaders might profitably
contemplate the Oyster: but such contemplation, to be profitable,
requires intelligence of a higher order than these protagonists of
folly appear to possess.
In justice to Darwin it should be remarked that he himself fully
realized, and carefully points out, the inconceivability of the
application of the Sexual Selection theory to the Mollusca. In
commenting on the beauty of colour and shape which many species
display, he remarks: “The colours do not appear in most cases, to be
of any use as a protection; they are probably the direct result, as in
the lowest classes, of _the nature of the tissues_[1]: the patterns and
the sculpture of the shell depending on the manner of growth.” Just
so: and this is surely the fundamental explanation of ornament, using
this term in its widest sense, everywhere in the Animal Kingdom. The
peculiarities and eccentricities of behaviour, which occur among the
higher groups, act as “aphrodisiacs” to hasten reproduction because
this confers an advantage, the earliest to produce offspring—so soon as
the conditions for their nurture are favourable—having the best chance
of survival. Premature sexual activity is checked by the death of the
offspring.
[1] Italics mine.
It has been contended that the hermaphrodite condition represents the
primitive mode of reproduction among the multicellular animals—that is
to say, all animals above the level of those whose bodies are composed
of but a single cell, or particle, of protoplasm—but this view is
probably erroneous, and the hermaphrodite state must be regarded as a
secondary condition, a later innovation.
More remarkable are the facts concerned with that singular form of
reproduction known as parthenogenesis, or the production of offspring
by virgin females. This is undoubtedly a degenerate sexual condition
occurring as a normal mode of reproduction, among the microscopic
“Rotifers,” _e.g._ the “Wheel-animalcule,” Crustacea, and Insects, and
in varying degrees of intensity.
The most familiar instances of Parthenogenesis are furnished by the
Hymenoptera, and notably by the Bees and the Aphides.
There are certain cases among the Rotifers where no males have ever
been found, and it is possible that they have become entirely
suppressed, but in every other case the periodical advent of males is
an absolute essential for the continuation of the race. Perhaps the
least degenerate of these types are the Bees, wherein we meet with
well-developed, highly-organized males and females, which, in their
sexual relationships, are perfectly normal. But in the fulfilment
of the mating instincts in these insects, a most amazing sequence
of events is revealed such as are without parallel in the rest of
the Animal Kingdom. The story has been charmingly told already by
Maeterlinck, in his delightful “Life of the Bee,” and it has been
told again by Tickner Edwardes, with less of poetry, perhaps, but
still fascinatingly: and it must be told again now, but in a condensed
fashion.
Briefly, a community of hive-bees harbours both male and female
individuals only for a very short space. During the greater part of the
year it consists only of a vast concourse of infertile females, the
daughters of one mother; the “queen” of the hive. The males of that
hive are the brothers, not the fathers, of the workers, as some have
supposed, and their sojourn there is brief. To gain a clear idea of the
facts in regard to the life-history of these insects it is necessary
to trace some of the incidents which lead up to the manner in which
the population of the hive is regulated, and its continuance ensured.
These may well begin with the time when the number of the inhabitants
consonant with the well-being of the hive has reached its limit. This
occurs during the early part of June, when the queen leaves the hive,
accompanied by several thousands of her daughters; they settle at
some distance from their late abode in a “swarm” for the purpose of
founding a new colony. Here we may leave them. The house just vacated
is, however, not entirely deserted. A few of the inhabitants, the
infertile sexless workers, degenerate females—degenerate so far as
the power of reproduction is concerned at any rate—are left behind,
and there remain also in their cradles a variable number of unhatched
queens, and drones or males. One of these potential queens and the
males now speedily emerge, and for a day or two remain within the
seclusion of the hive, feeding upon the honey stored in the combs.
The males are the first to leave, making daily excursions abroad in
the search for mates. They display in this a very leisurely behaviour,
rising late and not venturing out till the day is well aired. Returning
early in the afternoon with sharpened appetites, they feed to repletion
and soon fall asleep.
In about three days, however, the young queen ventures abroad, timidly
at first, to stretch her wings in the sunshine. She is preparing for
the great moment of her life, the nuptial flight. So far, though drones
may swarm on every side of her, no sign of recognition is given, nor do
the males evince any consciousness of her presence. She behaves warily
and demurely throughout. Her first excursions abroad are very brief;
they are not so much trial flights, apparently, as efforts to locate
the exact position of the hive in relation to the outer world. To this
end the flights are rapidly extended in ever-widening circles, till at
last, with lightning speed, she makes for the blue sky, to return to
the gloom of the hive almost immediately after. During all this time
the stimulus of sexual desire has been gathering force, and now, being
no longer controllable, she darts off, and up into the sky; almost
at once she is recognized by the swarms of males from neighbouring
hives, some thousands in number, which for days have been seeking this
event. Instantly they give eager chase, mounting after her higher and
ever higher. But as they ascend so their numbers decrease. Some, the
feeble, the ill-fed from impoverished hives, are speedily left behind;
many endure to the end, but only one secures the prize, and this great
moment of his life is also his last, for the fact of impregnation is
no sooner completed than Death claims him. He falls earthwards, as if
struck by lightning, and in his fall the intromittent organ is dragged
from his body, to be removed by the survivor of this mad flight, on her
descent.
She leaves a bride and returns a widow, filled with murderous
intentions. There are captive queens in the hive, and she can
tolerate no rivals. So soon as she has removed from her person the
embarrassing souvenir of her nuptial flight she makes for the Royal
cells. Accompanied by attendant workers she proceeds to tear off their
waxen coverings and put their occupants to death with a thrust of her
stiletto. No sooner is the work of execution over than the dead bodies
are seized by the workers and borne out of the hive. This awful task
is soon over, however, and henceforth for four or five long years she
remains a prisoner within the walls of her own palace. Craving neither
the air nor the light of the sun, she will die without once having
sipped the nectar from a flower. And during all this time, save during
the winter sleep, her sole duty is to produce sons and daughters. In
the prime of her maternity she may lay as many as three thousand eggs
a day. But strangely enough the number of eggs produced is determined
for her by the workers, who are the real rulers in this constitutional
state. By varying the amount and quality of the food they give her they
can increase or check the number of eggs produced; while even the sex
of the resultant larva is apparently also under their control.
During that brief, weird honeymoon in the clouds she received a store
of spermatozoa, the fertilizing male germs, sufficient for all the eggs
she can ever lay, and they may amount to nigh on a million. Incredible
as this may seem, their purpose is yet more so; for they are destined
to be expended solely in the production of female offspring doomed for
the most part to perpetual spinsterhood. One youngster in ten thousand
may attain to a higher state, may, if Fate wills, become a queen and
mother. And because of this need for mothers to carry on the race, this
extraordinary state of affairs has been brought about. All is under the
control of her daughters—the spinster-workers. As she proceeds on her
rounds of egg-laying an attendant crowd waits upon her, controlling
her actions by gentle caresses. As she passes from cell to cell, the
cradles of the young that are to be, she thrusts down her abdomen and
lays an egg in each. The cells destined to produce the workers are
the smallest, those for drones are larger, and those for queens are
largest of all, and the walls are formed of pure pollen, not of wax as
are those of the workers and drones. But it would seem that she never
lays an egg in any of the last named. The sight of a queen-cell rouses
her to fury. These cells, then, are filled by the workers, who remove
the requisite number of worker—eggs from the cells in which they were
laid and deposit them in the queen-cradles. The larvæ at hatching,
and for the first three days of life, differ in no wise from their
sisters around them. Their Royal state is determined solely by the food
which is administered to them. This consists of “bee-jelly,” which is
furnished in abundance: a white, shining liquid, regurgitated by the
ever-zealous nurse-bees. These superfed babies cease feeding at about
the fifth day, and each spins for herself a silken vestment in which to
undergo the pupal state. This done, the door of each cell is sealed up
with pollen. During the following sixteen days strange transformations
take place: the queen that is to be is taking shape. But the cradle now
becomes a prison, for at the end of the sixteenth day each of the four
or five young queens begins to clamour for release. But this cannot be,
for such as succeeded in emerging would immediately be slain by the
reigning queen. A small hole is bored through the roof of the cell, and
through this each is fed, and a close guard is kept night and day to
ensure that they shall not emerge till the moment is ripe. Soon each
captive begins to gnaw away the roof of her prison chamber, and as
rapidly more material is placed by her guards on the outer surface. Not
until the old queen leaves the hive with thousands of her daughters to
“swarm” and found a new colony will freedom be allowed; and then only
to one. The rest must remain till the new queen either also “swarms,”
or returns from her nuptial flight, and in this case all will be
slaughtered in their cramped quarters, unable to resist.
But what of the drone? He, as has already been mentioned, is reared
in a larger cradle than that of his sisters—save such as are destined
to be queens—and for the first three days of his life is fed on
“bee-milk” of a special kind and more generous quality than that of his
worker—sisters, the Cinderellas of the hive; but this generous diet
is diminished at the end of three days, when a mixture of honey and
pollen is given him. In about three weeks or rather more he emerges, a
great, lazy drone, and for a fortnight more he wanders about the hive
alternately soliciting bee-milk from his sisters and helping himself
to honey from the comb, and when full to repletion he seeks some
snug corner in which to sleep off his surfeit. In due time, however,
he ventures abroad, his hour is at hand. He takes his daily flights
abroad in search of a mate, returning home early in the afternoon for
his rations, being too indolent or too stupid to draw nectar from the
flowers for himself. Thus for many days he and his brothers disport
themselves in riotous living, till one or other of them attains the
end for which he was born; and after a few delirious moments drops
earthwards a mutilated corpse.
But so far only a part of the story of the drone’s life-history has
been told. Though the son of a queen, he has never had a father; and
should he ever attain to the dignity of fatherhood his posthumous
children are all daughters, most of whom die spinsters within six or
seven weeks of their birth, worn out by a life of ceaseless toil and
drudgery!
The queen, it will be remembered, cohabits with the male but once
in her life. The sperm-cells then received are stored in a special
receptacle and are released during the passage of the egg down the
oviduct. In this act of releasing the fertilizing germs a singular
economy is practised. In the case of most other creatures myriads of
sperm-cells are released for the fertilization of a single egg, and
of these but one can possibly attain its goal, the minute aperture or
“micropyle” which is the doorway to the germ liberated, in the form
of an egg, by the female. The rest die. In the case of the queen bee
but one of these precious sperm-cells is liberated at a time. Hence
her prolonged ability to produce fertilized eggs. But eggs destined
to produce males, or drones, are never thus fertilized: they are born
without the intervention of a father. A queen which has never mated
will lay only male-producing eggs. This is an astounding thing, but it
is true. No less remarkable is the fact that the sperm-cells should
survive in their encapsuled state for periods extending over several
years: it seems almost incredible, but it is nevertheless true.
One cannot suppose that the queen in coming to a drone cell
deliberately withholds the male germ as the egg passes down her
oviduct; some inhibitory factor preventing the release of the
sperm-cell must be brought into play which as yet we have not
discovered. This production of males from unfertilized eggs, or
“parthenogenesis” as it is called, is a common feature among the
hymenoptera, and some other groups of insects, and it occurs also among
other lowly creatures to be described later.
Having regard to the importance of the workers, a brief summary of
their life-history must be given. These, it has already been indicated,
are all, at any rate till three days old, potential queens. Their
development into, or degradation to, the lower grade is determined,
apparently, solely by the quality of the food, for the fact that
queens are reared only in specially constructed cells of large size
with walls of pollen instead of wax is explained by the larger size
of the queen and the need for a more porous, air-permeated cell-wall
on account of the longer time which must be spent in confinement.
The worker is certainly the most “intellectual” member of the hive,
but this superiority has been gained at a great price. Emerging from
the chrysalis skin at about three weeks from the time that the egg
from which she emerged was laid, she begins forthwith to gnaw her way
through the mass of wax and pollen which forms the door of her prison.
Rather, she eats her way through, for the material removed is swallowed
as it is detached, thus the young bee, as Mr. Tickner Edwardes remarks,
is caused to effect her own release by the promptings of her appetite.
Hunger-strikes in the bee community are unknown. Speedily the youngster
steps out, distinguishable from her elder sisters only by her weak,
grey-hued, flaccid appearance. Her first act on gaining freedom is to
groom herself down, after which she proceeds to explore the gloomy,
busy, crowded thoroughfares of the hive. A day or two is thus passed in
gathering strength. On the second appetite returns, and she proceeds
to help herself from the vats of honey and pollen bins scattered here
and there among the cradles of her sisters yet prisoners. But speedily
she is caught and thrust, so to speak, on to the treadmill of work
which is to know no cessation during her short span of life-some six
or seven weeks. Her first duties are those of nursemaid. Without
instruction, or previous experience, she begins to feed her younger
sisters and brothers yet in the larval stage. But besides, during her
first fortnight, before she is allowed to leave the hive she and her
sisters of the same age have to fulfil a variety of tasks. All the
indoor work of the house falls on these Cinderellas. Not only do they,
and they alone, feed the young, but they have to produce the wax and
build the combs and attend to the sanitary arrangements: “they are the
brewers of the honey and the keepers of the stores; they feed the queen
bee on her ceaseless rounds and give the drones, their brothers, their
daily rations of bee-milk”—what else these lazy creatures need they
take for themselves from the honey-vats. But this is not all. They have
to meet their older sisters returning from the fields and gardens laden
with nectar. This is regurgitated and transferred to the pouches of
the youngsters, by whom it is transformed into honey and stored in the
combs in the upper region of the hive. At the end of about a fortnight
these little drudges are allowed a brief respite, during the heat of
the day, to emerge into the outer air and gather ideas on the world
which is yet to be explored. Soon a measure of freedom is allowed, the
indoor work ceases, and each takes up the new and more agreeable task
of gathering pollen, and after a few days of this the more responsible
task of gathering nectar is undertaken, which is continued till death
ends one of the most crowded, surely, of existences. Such as are born
near “swarming-time” may have the good fortune to take part in the
exodus and the settling down in the new home, and some may taste yet
other moments of excitement, but they are moments only. The worker
bee knows no leisure for the improvement of her mind and morals. She
needs none, for she has neither: she is a creature of routine, a
living automaton apparently. Yet there are incidents in this wonderful
community which seem too complex to be merely the result of instinct
unaided, uninspired, by intelligence albeit of a nebulous kind.
The worker-bees, it has been remarked, are barren: their reproductive
organs are atrophied, and by the decree, not of the queen-mother of the
hive, nor of the males, but of their own caste. In spite of the fact
that they are incapable of producing offspring, they, and they alone,
determine who shall undertake this task; and they decree the fate that
awaits those thus appointed when they can no longer fulfil this purpose.
When the queen, waxing old, and waning in fecundity, lays fewer
and fewer eggs, and these only producing males, they take silent
note of the fact, and at the appointed time decree the death of
their Sovereign-mother. Yet they hesitate to lay violent hands on
her. She, as queen, claimed the right in her early youth to slay
her sister-queens, and sped them with a dagger-thrust; now her turn
comes to die. But it must be a bloodless death, carried out with due
ceremonial. So her daughters cluster about her, and in a mock embrace,
that tightens every moment, her breath is squeezed out of her body.
There are no State pensions for those who are past work, but a State
execution instead. This is vastly more economical, and it may yet
commend itself to some would-be social “reformers,” who will doubtless
contrive to make exceptions to the rule!
The execution of a queen is not an event of common occurrence; but
that of male members of the hive forms part of the ordinary routine,
though coming only within the larger cycle of the year. As the summer
wanes and the harvest of nectar grows perceptibly less, visions of a
possible famine, and its attendant horrors, seem to arise. So heads are
counted and occupations are scrutinized, when it is discovered that
the only members of the community who are contributing nothing to the
general well-being are the males, who are now but useless drains on
the hive. None of the neighbouring hives are now likely to send forth
a virgin queen to her nuptials, to which end each hive is obliged to
contribute—for no hive utilizes the services of its own drones; these
idle fellows, then, are “eating their heads off”—and males, too; perish
the thought! While they had anything to gain from him their motto
was “Feed the brute”; but now, on each, doom is pronounced. It must
be admitted that a live drone at the end of summer is one of life’s
failures. Notoriously unable to feed himself save upon the honey made
by his sisters, and having no function in life to perform save that of
mating, his very existence now is a damning witness against himself.
When the mother of the hive ceases to maintain the standard of
fertility set by her exacting daughters, she is put to death
stealthily, as if in an excess of devotion: she is smothered under
their embraces. Towards the drones now under sentence no such
consideration is to be shown. When the word goes forth, the slaughter
begins, and it gathers in ferocity. It begins in a massacre of the
innocents—every helpless larval drone is ruthlessly dragged from
its cot and thrown out of the hive to die: there is now no crime in
infanticide, nor in the most gruesome massacre that is presently to
follow. The drones, all unsuspecting, are to be tolerated a brief spell
longer. The cool, calculating spirit of these unsexed ones seems to
realize that there is even yet a remote possibility that the services
of these doomed ones may be wanted. No sooner, however, does it become
clear that this chance is past, than the decree of death is made
absolute, and the poor drones are suddenly and viciously attacked by
half a dozen frenzied spinsters at once. Each tries to bite through
the base of the victim’s wings, and succeeding in this, he is speedily
pushed towards the door of the hive and out into the open, whence
return is impossible, so that nothing is left but death by starvation.
Some of the victims will escape in the _mêlée_, but only for a brief
season. Such as find their way, unmaimed, to the open air, are still
faced by inevitable death. To remain out is to die of starvation or
cold, to return is to fall a prey to the now infuriated guards, who,
strongly reinforced, stand at the doorway of the hive to intercept
and dispatch these unlucky fugitives. It will be remarked that these
executioners make no use of their stings; these they might be unable to
withdraw from their victim’s body, in which case they, too, would die.
But there is no need to run this risk, for the males, their brothers,
whom they so cheerfully slay, are unarmed; they may be attacked without
risk. The dreadful work, however, is soon over, and the survivors, the
queen and her daughters, have the house to themselves to make the final
preparations for the winter sleep, which is apparently undisturbed by
qualms of conscience.
There are certain structural differences distinguishing the three types
in such a hive—the queen, the drone and the worker—which must now be
referred to. The queen is larger than the worker; she has a larger
and longer abdomen, a longer and much-curved sting, and her eyes have
fewer facets. Only vestiges remain of the wax-secreting organs, and no
trace is to be found of the wonderful pollen-baskets which perform so
important a function in the worker; and finally, her instincts are of a
very different kind.
The “pollen-basket” of the worker is a strange contrivance. The pollen
is mainly collected by the hairs which clothe the under surface of
the body, from which it is scraped by special brushes of hairs which
clothe the inner surface of the “metatarsus “—the big, flat joint to
which are attached a series of small triangular joints, the last of
which bears the claws. When the brushes are “clogged up,” the legs are
crossed and the pollen is combed out by specially stiff hairs on the
“tibia”—the joint immediately above the metatarsus—and the bolus thus
formed is then transferred to the outer surface of the tibia, which is
trough-shaped, forming the “corbiculum,” or pollen-basket. The next,
or middle, pair of legs are then employed to ram the pollen well into
the basket, for safe conveyance to the hive. On arrival at the combs,
the bee pushes its hind-legs into a cell, or “pollen-tub,” and with a
special spur dislodges the pellet of pollen and lets it fall into the
tub. These are complex movements, performed without instruction and, we
must suppose, without any intelligent conception of their purpose.
The drone is larger than either queen or worker, and has enormous
eyes, which meet one another over the top of the head; he has no
wax-secreting organs, no pollen-basket, no sting. His antennæ are
longer, his hum is deeper, his sole function is to fertilize a queen,
and this done, he promptly dies. Failing in his first flight, he may
make yet other ventures, but the chances are that he will die without
attaining the only purpose for which he exists.
The fact that he lives for some days in the hive with the queen,
before her nuptial flight, apparently unaware of her presence, would
seem to indicate some special “trigger” for the release of the sexual
instincts. But it must be remembered that he does not attain to
maturity until after his first flight, and this it is, probably, which
arouses the mate-hunger. More than this, however, it is probable that
coitus is possible only when on the wing, when the air-sacs become
inflated, and exert pressure on the genital organs. How he recognizes
the queen when on her wild flight heavenwards is unknown: possibly by
scent, but more probably by the very different vibrative note of her
wings, that of the male being much stronger and deeper. His continued
return to the hive is a proof of his failure to justify his existence,
for no drone ever experienced Love’s embrace and lived to tell the
tale: hence, when the time comes, he is slain without compunction.
These differences between the fully-developed male and female present
nothing very striking; but how are the singular peculiarities of
structure and instinct in the “workers” to be accounted for? They are
present in neither queen nor drone, yet by them they are transmitted
to their offspring from one generation to another! It is true that
every worker, for a time, is a potential queen, and every queen, but
for the grace of Chance, might have been a worker. All depends on the
food. It is remarkable, but apparently the fact, that a more generous
diet, or, rather, a more stimulating diet, should so profoundly modify
the organism, but, it is to be noted, this sleight-of-hand is only
successfully practised on a larva during its first three days of
existence. Thus the royal bee jelly stimulates the growth of the sexual
organs and inhibits the development of the structures peculiar to the
worker—the basket, and pollen-hairs, and so on. These structures are
not made by the food; they are simply nourished or inhibited, as the
case may be. Nevertheless, one cannot help being mystified by the fact
that the mere difference in the quality of the food, or, rather, in the
chemical constituents thereof, should cause the inhibition, or, rather,
the suppression, of relatively complex structures like the corbiculum
and the reduction of the number of the facets of the eye. To say that
the structures inhibited, in the case of the queen, are just those
which will be of no service when in her royal state, is by no means to
explain the mystery. And what is true of the physical side is no less
true of the psychical, for with this change of diet the behaviour of
the insect, throughout its whole life, is most profoundly changed. If
the pollen-basket is wanting, no less so are the instinctive actions
associated with its use; if the genital organs are atrophied, so also
are the instinctive acts associated therewith. This nexus between
instinct and structure is not to be lost sight of.
How—and the question has often been asked—are the experiences of the
infertile females, the workers, transmitted to the germ-plasm? For the
workers, it has been contended, being sterile, are incapable of handing
on such acquirements: this is so. These workers hold the same position
in regard to the species that structures essential to well-being hold
in regard to the individual. These last are not under the control of
the individual, but are determined by a plus or minus quality in its
germ-plasm. The worker-bees are products of the germ-plasm, committed
to the care of the queens. Any strain, so to speak, of that germ-plasm
which gives rise to defective workers brings about its own extinction,
or elimination, sooner or later. Any strain of germ-plasm which
contains, so to speak, a spark of that quality which in the individual
is expressed by intelligent behaviour, will gain advantages in the
struggle for existence.
The complex, the extraordinarily complex, behaviour of the worker-bees
on any interpretation is still mysterious. This interpretation can be
tested only by a reference to the life-history of other social-bees
which have attained to a less complexity. This shows us that the
sterile worker is not to be regarded as a newly-evolved type so much as
an arrested stage of a more complete ancestral condition, and the fact
that the worker is potentially a queen is further evidence of this.
A clue to many of the more puzzling features presented by the domestic
economy of the Hive-bee may be obtained by a study of the life-history
of other species of social-bees which have not attained to so high a
degree of specialization. The Bumble-bees afford illustrations of the
stages through which _Apis mellifica_, the Hive-bee, must have passed.
In the stone Bumble-bee (_Bombus lapidarius_), a queen, who has passed
the winter in blissful sleep, will lay the foundation for a new colony
on some bright May morning by collecting a small quantity of moss. This
done, she starts forth to gather pollen, with which, under cover of the
moss, she forms a waxen cell, mixing the newly-gathered pollen with the
wax so mysteriously formed within her body, as in the case of Hive-bees
of the worker type. Slowly and laboriously this waxen cradle grows.
Fashioned like a globe, its inner surface is lined with pollen soaked
in honey, and with the last pellet of this a number of eggs are laid
arid the nursery is sealed up. By the time these labours are completed
the queen is worn out; she therefore rests awhile, clinging to the
outer wall of this cunningly-wrought cradle. After a few days’ rest she
adds another and commonly yet a third cell to the first, joining each
to the other with wax. But before the third cradle is finished the eggs
in the first have hatched. The youngsters will have consumed the layer
of honey-soaked pollen placed there for this purpose. They therefore
require feeding, and thus the labours of this very industrious queen
are still further increased. Divining the needs of her imprisoned
first-born, she bites a small hole through the nursery wall and pours
in a quantity of honey for their sustenance. In due time they are
“full-fed,” and each spins for itself a silken vestment wherein to
undergo its transformation into a worker-bee. The careful mother,
during this period of transition, now scrapes away an opening through
which the young bees may creep when they awake. This event takes place
in the course of a few days, when her work is materially lightened, for
these newly-hatched workers at once take over the duties of building
nurseries and feeding the further batches of young which, for a time,
follow one another in quick succession. The queen, indeed, has now
nothing else to do but to lay eggs in the nurseries as they are ready.
So far all the children born to her are daughters. The earliest-born,
it is to be noted, were “workers”; those which follow and are tended by
the workers are also females, and supplement their mother’s labours by
producing fertile eggs, though they have never even seen the male of
their own species. Thus, if the queen-mother die her virgin daughters
carry on the colony. But it sometimes happens that she may have left
no descendants capable, for the time, of laying fertile eggs. In this
case, if there be larvæ still in the nursery, the workers feed them
assiduously as if in the hope that some may prove fertile. But if
there be no infants to be fed they apparently abandon work, become
despondent, and spend the greater part of their time sitting at home
by the empty cradles, till at last death comes to their rescue and the
colony is extinct.
Much that baffles one in the history of the Hive-bee becomes clear in
the light of the facts revealed by the life-story of the Bumble-bee.
In the first place it will be remembered her first eggs produced only
workers, which appeared at a time when her energies were severely
strained, and their food allowance was no more than barely sufficient
to sustain life. The females which appeared later produced fertile
eggs, having been more abundantly fed by their infertile elder sisters.
The number of fertile females which appear at this stage of the colony
seems again to be regulated by the abundance of food, which varies
in amount with fine, or cold, weather. Even among the worker broods
fertile females may appear. They owe their fertility apparently to good
luck, which afforded them the opportunity of securing more food than
their sisters. The birth of young from females about whose virginity
there can be no question is certainly remarkable, but it would seem
that this parthenogenetic state is one of limited endurance, for
towards the end of summer males appear, and these mating with some of
the later-born females, lead again to the appearance of a queen, who,
being fertilized, alone survives the winter to carry on the race with
the succeeding summer.
Thus, then, the mysterious existence of the workers among the
Hive-bees, displaying structural peculiarities and instincts so
different from those of the queen-mother, is explained. For the queen,
in this case, is evidently the product of a more intensified, more
perfected, social system, relieved, from the first, of the labours
of building and the care of her offspring, duties which the queen
Bumble-bee has at first to perform for herself, because all her
children die at the end of the summer. Among Hive-bees fertile workers
also occasionally occur; they are probably bees which in their larval
state received a more than usually abundant supply of food, or food
approximating to the “bee jelly” which produces young queens. The
difference, then, between the individuals of a colony of Hive-bees and
one of Bumble-bees lies in the greater abundance of fertile workers
and in the fact that the queen of the Hive-bees is relieved of all
work from the first, and so is enabled to devote her whole energies to
the duties of reproduction. She is the descendant of a race of queens
which in earlier times, like the Bumble-bee queen, had to perform
the duties now relegated to her daughters, who inherit not only her
house-building and child-nurturing instincts, but also her potentiality
for child-bearing, though this potentiality is commonly inhibited
by the starvation of the reproductive activities. Selection secures
survival of this state of affairs by the elimination of any tendency
to lose any of these qualities on the part of the queen. The workers
of the Hive-bee, in short, have not evolved their peculiarities of
structure and instinct by some mysterious process of natural selection
confined to the workers individually, for these, being infertile, could
not transmit any of their inherent qualities or tendencies to variation
in the direction of more efficient workers. On the contrary, all that
they possess they inherit from the queen-mother, who transmits to her
offspring the qualities and characteristics her forebears in the female
line possessed in their own person.