Chapter V. Permutations of Sex
Mason Bee Nest in a shell
The sex of the egg is optional. The choice rests with the mother, who is guided by considerations of space and, according to the accommodation at her disposal, which is frequently fortuitous and incapable of modification, places a female in this cell and a male in that, so that both may have a dwelling of a size suited to their unequal development. This is the unimpeachable evidence of the numerous and varied facts which I have set forth. People unfamiliar with insect anatomy — the public for whom I write — would probably give the following explanation of this marvellous prerogative of the Bee: the mother has at her disposal a certain number of eggs, some of which are irrevocably female and the others irrevocably male: she is able to pick out of either group the one which she wants at the actual moment; and her choice is decided by the holding capacity of the cell that has to be stocked. Everything would then be limited to a judicious selection from the heap of eggs.
Should this idea occur to him, the reader must hasten to reject it. Nothing could be more false, as the merest reference to anatomy will show. The female reproductive apparatus of the Hymenoptera consists generally of six ovarian tubes, something like glove-fingers, divided into bunches of three and ending in a common canal, the oviduct, which carries the eggs outside. Each of these glove-fingers is fairly wide at the base, but tapers sharply towards the tip, which is closed. It contains, arranged in a row, one after the other, like beads on a string, a certain number of eggs, five or six for instance, of which the lower ones are more or less developed, the middle ones half-way towards maturity, and the upper ones very rudimentary. Every stage of evolution is here represented, distributed regularly from bottom to top, from the verge of maturity to the vague outlines of the embryo. The sheath clasps its string of ovules so closely that any inversion of the order is impossible. Besides, an inversion would result in a gross absurdity: the replacing of a riper egg by another in an earlier stage of development.
Therefore, in each ovarian tube, in each glove-finger, the emergence of the eggs occurs according to the order governing their arrangement in the common sheath; and any other sequence is absolutely impossible. Moreover, at the nesting period, the six ovarian sheaths, one by one and each in its turn, have at their base an egg which in a very short time swells enormously. Some hours or even a day before the laying, that egg by itself represents or even exceeds in bulk the whole of the ovigenous apparatus. This is the egg which is on the point of being laid. It is about to descend into the oviduct, in its proper order, at its proper time; and the mother has no power to make another take its place. It is this egg, necessarily this egg and no other, that will presently be laid upon the provisions, whether these be a mess of honey or a live prey; it alone is ripe, it alone is at the entrance to the oviduct; none of the others, since they are farther back in the row and not at the right stage of development, can be substituted at this crisis. Its birth is inevitable.
What will it yield, a male or a female? No lodging has been prepared, no food collected for it; and yet both food and lodging have to be in keeping with the sex that will proceed from it. And here is a much more puzzling condition: the sex of that egg, whose advent is predestined, has to correspond with the space which the mother happens to have found for a cell. There is therefore no room for hesitation, strange though the statement may appear: the egg, as it descends from its ovarian tube, has no determined sex. It is perhaps during the few hours of its rapid development at the base of its ovarian sheath, it is perhaps on its passage through the oviduct that it receives, at the mother's pleasure, the final impress that will produce, to match the cradle which it has to fill, either a female or a male.
Thereupon the following question presents itself. Let us admit that, when the normal conditions remain, a laying would have yielded m females and n males. Then, if my conclusions are correct, it must be in the mother's power, when the conditions are different, to take from the m group and increase the n group to the same extent; it must be possible for her laying to be represented as m-1, m-2, m-3, etc. females and by n+1, n+2, n+3, etc. males, the sum of m+n remaining constant, but one of the sexes being partly permuted into the other. The ultimate conclusion even cannot be disregarded: we must admit a set of eggs represented by m-m, or zero, females and of n+m males, one of the sexes being completely replaced by the other. Conversely, it must be possible for the feminine series to be augmented from the masculine series to the extent of absorbing it entirely. It was to solve this question and some others connected with it that I undertook, for the second time, to rear the Three-horned Osmia in my study.
The problem on this occasion is a more delicate one; but I am also better-equipped. My apparatus consists of two small, closed packing- cases, with the front side of each pierced with forty holes, in which I can insert my glass tubes and keep them in a horizontal position. I thus obtain for the Bees the darkness and mystery which suit their work and for myself the power of withdrawing from my hive, at any time, any tube that I wish, with the Osmia inside, so as to carry it to the light and follow, if need be with the aid of the lens, the operations of the busy worker. My investigations, however frequent and minute, in no way hinder the peaceable Bee, who remains absorbed in her maternal duties.
I mark a plentiful number of my guests with a variety of dots on the thorax, which enables me to follow any one Osmia from the beginning to the end of her laying. The tubes and their respective holes are numbered; a list, always lying open on my desk, enables me to note from day to day, sometimes from hour to hour, what happens in each tube and particularly the actions of the Osmiae whose backs bear distinguishing marks. As soon as one tube is filled, I replace it by another. Moreover, I have scattered in front of either hive a few handfuls of empty Snail-shells, specially chosen for the object which I have in view. Reasons which I will explain later led me to prefer the shells of Helix caespitum. Each of the shells, as and when stocked, received the date of the laying and the alphabetical sign corresponding with the Osmia to whom it belonged. In this way, I spent five or six weeks in continual observation. To succeed in an enquiry, the first and foremost condition is patience. This condition I fulfilled; and it was rewarded with the success which I was justified in expecting.
The tubes employed are of two kinds. The first, which are cylindrical and of the same width throughout, will be of use for confirming the facts observed in the first year of my experiments in indoor rearing. The others, the majority, consist of two cylinders which are of very different diameters, set end to end. The front cylinder, the one which projects a little way outside the hive and forms the entrance-hole, varies in width between 8 and 12 millimetres (1). The second, the back one, contained entirely within my packing-case, is closed at its far end and is 5 to 6 millimetres (2) in diameter. Each of the two parts of the double-galleried tunnel, one narrow and one wide, measures at most a decimetre (3) in length. I thought it advisable to have these short tubes, as the Osmia is thus compelled to select different lodgings, each of them being insufficient in itself to accommodate the total laying. In this way I shall obtain a greater variety in the distribution of the sexes. Lastly, at the mouth of each tube, which projects slightly outside the case, there is a little paper tongue, forming a sort of perch on which the Osmia alights on her arrival and giving easy access to the house. With these facilities, the swarm colonized fifty-two double-galleried tubes, thirty-seven cylindrical tubes, seventy-eight Snail-shells and a few old nests of the Mason-bee of the Shrubs. From this rich mine of material I will take what I want to prove my case.
Every series, even when incomplete, begins with females and ends with males. To this rule I have not yet found an exception, at least in galleries of normal diameter. In each new abode, the mother busies herself first of all with the more important sex. Bearing this point in mind, would it be possible for me, by manoeuvring, to obtain an inversion of this order and make the laying begin with males? I think so, from the results already ascertained and the irresistible conclusions to be drawn from them. The double-galleried tubes are installed in order to put my conjectures to the proof.
The back gallery, 5 or 6 millimetres (4) wide, is too narrow to serve as a lodging for normally developed females. If, therefore, the Osmia, who is very economical of her space, wishes to occupy them, she will be obliged to establish males there. And her laying must necessarily begin here, because this corner is the rear-most part of the tube. The foremost gallery is wide, with an entrance-door on the front of the hive. Here, finding the conditions to which she is accustomed, the mother will go on with her laying in the order which she prefers.
Let us now see what has happened. Of the fifty-two double galleried tubes, about a third did not have their narrow passage colonized. The Osmia closed its aperture communicating with the large passage; and the latter alone received the eggs. This waste of space was inevitable. The female Osmiae, though nearly always larger than the males, present marked differences among one another: some are bigger, some are smaller. I had to adjust the width of the narrow galleries to Bees of average dimensions. It may happen therefore that a gallery is too small to admit the large-sized mothers to whom chance allots it. When the Osmia is unable to enter the tube, obviously she will not colonize it. She then closes the entrance to this space which she cannot use and does her laying beyond it, in the wide tube. Had I tried to avoid these useless apparatus by choosing tubes of larger calibre, I should have encountered another drawback: the medium-sized mothers, finding themselves almost comfortable, would have decided to lodge females there. I had to be prepared for it: as each mother selected her house at will and as I was unable to interfere in her choice, a narrow tube would be colonized or not, according as the Osmia who owned it was or was not able to make her way inside.
There remain some forty pairs of tubes with both galleries colonized. In these there are two things to take into consideration. The narrow rear tubes of 5 or 5 1/2 millimetres (5) — and these are the most numerous — contain males and males only, but in short series, between one and five. The mother is here so much hampered in her work that they are rarely occupied from end to end; the Osmia seems in a hurry to leave them and to go and colonize the front tube, whose ample space will leave her the liberty of movement necessary for her operations. The other rear tubes, the minority, whose diameter is about 6 millimetres (6), contain sometimes only females and sometimes females at the back and males towards the opening. One can see that a tube a trifle wider and a mother slightly smaller would account for this difference in the results. Nevertheless, as the necessary space for a female is barely provided in this case, we see that the mother avoids as far as she can a two-sex arrangement beginning with males and that she adopts it only in the last extremity. Finally, whatever the contents of the small tube may be, those of the large one, following upon it, never vary and consist of females at the back and males in front.
Though incomplete, because of circumstances very difficult to control, the result of the experiment is none the less very striking. Twenty-five apparatus contain only males in their narrow gallery, in numbers varying from a minimum of one to a maximum of five. After these comes the colony of the large gallery, beginning with females and ending with males. And the layings in these apparatus do not always belong to late summer or even to the intermediate period: a few small tubes contain the earliest eggs of the Osmiae. A couple of Osmiae, more forward than the others, set to work on the 23rd of April. Both of them started their laying by placing males in the narrow tubes. The meagre supply of provisions was enough in itself to show the sex, which proved later to be in accordance with my anticipations. We see then that, by my artifices, the whole swarm starts with the converse of the normal order. This inversion is continued, at no matter what period, from the beginning to the end of the operations. The series which, according to rule, would begin with females now begins with males. Once the larger gallery is reached, the laying is pursued in the usual order.
We have advanced one step and that no small one: we have seen that the Osmia, when circumstances require it, is capable of reversing the sequence of the sexes. Would it be possible, provided that the tube were long enough, to obtain a complete inversion, in which the entire series of the males should occupy the narrow gallery at the back and the entire series of the females the roomy gallery in front? I think not; and I will tell you why.
Long and narrow cylinders are by no means to the Osmia's taste, not because of their narrowness but because of their length. Remember that for each load of honey brought the worker is obliged to move backwards twice. She enters, head first, to begin by disgorging the honey-syrup from her crop. Unable to turn in a passage which she blocks entirely, she goes out backwards, crawling rather than walking, a laborious performance on the polished surface of the glass and a performance which, with any other surface, would still be very awkward, as the wings are bound to rub against the wall with their free end and are liable to get rumpled or bent. She goes out backwards, reaches the outside, turns round and goes in again, but this time the opposite way, so as to brush off the load of pollen from her abdomen on to the heap. If the gallery is at all long, this crawling backwards becomes troublesome after a time; and the Osmia soon abandons a passage that is too small to allow of free movement. I have said that the narrow tubes of my apparatus are, for the most part, only very incompletely colonized. The Bee, after lodging a small number of males in them, hastens to leave them. In the wide front gallery, she can stay where she is and still be able to turn round easily for her different manipulations; she will avoid those two long journeys backwards, which are so exhausting and so bad for her wings.
Another reason no doubt prompts her not to make too great a use of the narrow passage, in which she would establish males, followed by females in the part where the gallery widens. The males have to leave their cells a couple of weeks or more before the females. If they occupy the back of the house, they will die prisoners or else they will overturn everything on their way out. This risk is avoided by the order which the Osmia adopts.
In my tubes with their unusual arrangement, the mother might well find the dilemma perplexing: there is the narrowness of the space at her disposal and there is the emergence later on. In the narrow tubes, the width is insufficient for the females; on the other hand, if she lodges males there, they are liable to perish, since they will be prevented from issuing at the proper moment. This would perhaps explain the mother's hesitation and her obstinacy in settling females in some of my apparatus which looked as if they could suit none but males.
A suspicion occurs to me, a suspicion aroused by my attentive examination of the narrow tubes. All, whatever the number of their inmates, are carefully plugged at the opening, just as separate tubes would be. It might therefore be the case that the narrow gallery at the back was looked upon by the Osmia not as the prolongation of the large front gallery, but as an independent tube. The facility with which the worker turns as soon as she reaches the wide tube, her liberty of action, which is now as great as in a doorway communicating with the outer air, might well be misleading and cause the Osmia to treat the narrow passage at the back as though the wide passage in front did not exist. This would account for the placing of the female in the large tube above the males in the small tube, an arrangement contrary to her custom.
I will not undertake to decide whether the mother really appreciates the danger of my snares, or whether she makes a mistake in considering only the space at her disposal and beginning with males. At any rate, I perceive in her a tendency to deviate as little as possible from the order which safeguards the emergence of the two sexes. This tendency is demonstrated by her repugnance to colonizing my narrow tubes with long series of males. However, so far as we are concerned, it does not matter much what passes at such times in the Osmia's little brain. Enough for us to know that she dislikes narrow and long tubes, not because they are narrow, but because they are at the same time long.
And, in fact, she does very well with a short tube of the same diameter. Such are the cells in the old nests of the Mason-bee of the Shrubs and the empty shells of the Garden Snail. With the short tube, the two disadvantages of the long tube are avoided. She has very little of that crawling backwards to do when she has a Snail-shell for the home of her eggs and scarcely any when the home is the cell of the Mason-bee. Moreover, as the stack of cocoons numbers two or three at most, the deliverance will be exempt from the difficulties attached to a long series. To persuade the Osmia to nidify in a single tube long enough to receive the whole of her laying and at the same time narrow enough to leave her only just the possibility of admittance appears to me a project without the slightest chance of success: the Bee would stubbornly refuse such a dwelling or would content herself with entrusting only a very small portion of her eggs to it. On the other hand, with narrow but short cavities, success, without being easy, seems to me at least quite possible. Guided by these considerations, I embarked upon the most arduous part of my problem: to obtain the complete or almost complete permutation of one sex with the other; to produce a laying consisting only of males by offering the mother a series of lodgings suited only to males.
Let us in the first place consult the old nests of the Mason-bee of the Shrubs. I have said that these mortar spheroids, pierced all over with little cylindrical cavities, are adopted pretty eagerly by the Three-horned Osmia, who colonizes them before my eyes with females in the deep cells and males in the shallow cells. That is how things go when the old nest remains in its natural state. With a grater, however, I scrape the outside of another nest so as to reduce the depth of the cavities to some ten millimetres (7). This leaves in each cell just room for one cocoon, surmounted by the closing stopper. Of the fourteen cavities in the nests, I leave two intact, measuring fifteen millimetres in depth (8). Nothing could be more striking than the result of this experiment, made in the first year of my home rearing. The twelve cavities whose depth had been reduced all received males; the two cavities left untouched received females.
A year passes and I repeat the experiment with a nest of fifteen cells; but this time all the cells are reduced to the minimum depth with the grater. Well, the fifteen cells, from first to last, are occupied by males. It must be quite understood that, in each case, all the offspring belonged to one mother, marked with her distinguishing spot and kept in sight as long as her laying lasted. He would indeed be difficult to please who refused to bow before the results of these two experiments. If, however, he is not yet convinced, here is something to remove his last doubts.
The Three-horned Osmia often settles her family in old shells, especially those of the Common Snail (Helix aspersa), who is so common under the stone-heaps and in the crevices of the little unmortared walls that support our terraces. In this species, the spiral is wide open, so that the Osmia, penetrating as far down as the helical passage permits, finds, immediately above the point which is too narrow to pass, the space necessary for the cell of a female. This cell is succeeded by others, wider still, always for females, arranged in a line in the same way as in a straight tube. In the last whorl of the spiral, the diameter would be too great for a single row. Then longitudinal partitions are added to the transverse partitions, the whole resulting in cells of unequal dimensions in which males predominate, mixed with a few females in the lower storeys. The sequence of the sexes is therefore what it would be in a straight tube and especially in a tube with a wide bore, where the partitioning is complicated by subdivisions on the same level. A single Snail-shell contains room for six or eight cells. A large, rough earthen stopper finishes the nest at the entrance to the shell.
As a dwelling of this sort could show us nothing new, I chose for my swarm the Garden Snail (Helix caespitum), whose shell, shaped like a small, swollen Ammonite, widens by slow degrees, the diameter of the usable portion, right up to the mouth, being hardly greater than that required by a male Osmia-cocoon. Moreover, the widest part, in which a female might find room, has to receive a thick stopping-plug, below which there will often be a free space. Under all these conditions, the house will hardly suit any but males arranged one after the other.
The collection of shells placed at the foot of each hive includes specimens of different sizes. The smallest are 18 millimetres (9) in diameter and the largest 24 millimetres (10). There is room for two cocoons, or three at most, according to their dimensions.
Now these shells were used by my visitors without any hesitation, perhaps even with more eagerness than the glass tubes, whose slippery sides might easily be a little annoying to the Bee. Some of them were occupied on the first few days of the laying; and the Osmia who had started with a home of this sort would pass next to a second Snail-shell, in the immediate neighbourhood of the first, to a third, a fourth and others still, always close to one another, until her ovaries were emptied. The whole family of one mother would thus be lodged in Snail-shells which were duly marked with the date of the laying and a description of the worker. The faithful adherents of the Snail-shell were in the minority. The greater number left the tubes to come to the shells and then went back from the shells to the tubes. All, after filling the spiral staircase with two or three cells, closed the house with a thick earthen stopper on a level with the opening. It was a long and troublesome task, in which the Osmia displayed all her patience as a mother and all her talents as a plasterer. There were even some who, scrupulous to excess, carefully cemented the umbilicus, a hole which seemed to inspire them with distrust as being able to give access to the interior of the dwelling. It was a dangerous-looking cavity, which for the greater safety of the family it was prudent to block up.
When the pupae are sufficiently matured, I proceed to examine these elegant abodes. The contents fill me with joy: they fulfil my anticipations to the letter. The great, the very great majority of the cocoons turn out to be males; here and there, in the bigger cells, a few rare females appear. The smallness of the space has almost done away with the sixty-eight Snail-shells colonized. But, of this total number, I must use only those series which received an entire laying and were occupied by the same Osmia from the beginning to the end of the egg-season. Here are a few examples, taken from among the most conclusive.
>From the 6th of May, when she started operations, to the 25th of May, the date at which her laying ceased, the Osmia occupied seven Snail- shells in succession. Her family consists of fourteen cocoons, a number very near the average; and, of these fourteen cocoons, twelve belong to males and only two to females. These occupy the seventh and thirteenth places in chronological order.
Another, between the 9th and 27th of May, stocked six Snail-shells with a family of thirteen, including ten males and three females. The places occupied by the latter in the series were numbers 3, 4 and 5.
A third, between the 2nd and 29th of May, colonized eleven Snail-shells, a prodigious task. This industrious one was also exceedingly prolific. She supplied me with a family of twenty-six, the largest which I have ever obtained from one Osmia. Well, this abnormal progeny consisted of twenty-five males and one female, one alone, occupying place 17.
There is no need to go on, after this magnificent example, especially as the other series would all, without exception, give us the same result. Two facts are immediately obvious. The Osmia is able to reverse the order of her laying and to start with a more or less long series of males before producing any females. In the first case, the first female appears as number 7; in the third, as number 17. There is something better still; and this is the proposition which I was particularly anxious to prove: the female sex can be permuted with the male sex and can be permuted to the point of disappearing altogether. We see this especially in the third case, where the presence of a solitary female in a family of twenty-six is due to the somewhat larger diameter of the corresponding Snail-shell and also, no doubt, to some mistake on the mother's part, for the female cocoon, in a series of two, occupies the upper storey, the one next to the orifice, an arrangement which the Osmia appears to me to dislike.
This result throws so much light on one of the darkest corners of biology that I must attempt to corroborate it by means of even more conclusive experiments. I propose next year to give the Osmiae nothing but Snail-shells for a lodging, picked out one by one, and rigorously to deprive the swarm of any other retreat in which the laying could be effected. Under these conditions, I ought to obtain nothing but males, or nearly, for the whole swarm.
There would still remain the inverse permutation: to obtain only females and no males, or very few. The first permutation makes the second seem very probable, although I cannot as yet conceive a means of realizing it. The only condition which I can regulate is the dimensions of the home. When the rooms are small, the males abound and the females tend to disappear. With generous quarters, the converse would not take place. I should obtain females and afterwards an equal number of males, confined in small cells which, in case of need, would be bounded by numerous partitions. The factor of space does not enter into the question here. What artifice can we then employ to provoke this second permutation? So far, I can think of nothing that is worth attempting.
It is time to conclude. Leading a retired life, in the solitude of a village, having quite enough to do with patiently and obscurely ploughing my humble furrow, I know little about modern scientific views. In my young days I had a passionate longing for books and found it difficult to procure them; to-day, when I could almost have them if I wanted, I am ceasing to wish for them. It is what usually happens as life goes on. I do not therefore know what may have been done in the direction whither this study of the sexes has led us. If I am stating propositions that are really new or at least more comprehensive than the propositions already known, my words will perhaps sound heretical. No matter: as a simple translator of facts, I do not hesitate to make my statement, being fully persuaded that time will turn my heresy into orthodoxy. I will therefore recapitulate my conclusions.
Bees lay their eggs in series of first females and then males, when the two sexes are of different sizes and demand an unequal quantity of nourishment. When the two sexes are alike in size, the same sequence may occur, but less regularly.
This dual arrangement disappears when the place chosen for the nest is not large enough to contain the entire laying. We then see broken layings, beginning with females and ending with males.
The egg, as it issues from the ovary, has not yet a fixed sex. The final impress that produces the sex is given at the moment of laying or a little before.
So as to be able to give each larva the amount of space and food that suits it according as it is male or female, the mother can choose the sex of the egg which she is about to lay. To meet the conditions of the building, which is often the work of another or else a natural retreat that admits of little or no alteration, she lays either a male egg or a female egg as she pleases. The distribution of the sexes depends upon herself. Should circumstances require it, the order of the laying can be reversed and begin with males; lastly, the entire laying can contain only one sex.
The same privilege is possessed by the predatory Hymenoptera, the Wasps, at least by those in whom the two sexes are of a different size and consequently require an amount of nourishment that is larger in the one case than in the other. The mother must know the sex of the egg which she is going to lay; she must be able to choose the sex of that egg so that each larva may obtain its proper portion of food.
Generally speaking, when the sexes are of different sizes, every insect that collects food and prepares or selects a dwelling for its offspring must be able to choose the sex of the egg in order to satisfy without mistake the conditions imposed upon it.
The question remains how this optional assessment of the sexes is effected. I know absolutely nothing about it. If I should ever learn anything about this delicate point, I shall owe it to some happy chance for which I must wait, or rather watch, patiently. Towards the end of my investigations, I heard of a German theory which relates to the Hive-bee and comes from Dzierzon, the apiarist(11). If I understand it aright, according to the very incomplete documents which I have before me, the egg, as it issues from the ovary, is said already to possess a sex, which is always the same; it is originally male; and it becomes female by fertilization. The males are supposed to proceed from non-fertilized eggs, the females from fertilized eggs. The Queen-bee would thus lay female eggs or male eggs according as she fertilized them or not while they were passing into her oviduct.
Coming from Germany, this theory cannot but inspire me with profound distrust. As it has been given acceptance, with rash precipitancy, in standard works, I will overcome my reluctance to devoting my attention to Teutonic ideas and will submit it not to the test of argument, which can always be met by an opposite argument, but to the unanswerable test of facts.
For this optional fertilization, determining the sex, the mother's organism requires a seminal reservoir which distils its drop of sperm upon the egg contained in the oviduct and thus gives it a feminine character, or else leaves it its original character, the male character, by refusing it that baptism. This reservoir exists in the Hive-bee. Do we find a similar organ in the other Hymenoptera, whether honey-gatherers or hunters? The anatomical treatises are either silent on this point or, without further enquiry, apply to the order as a whole the data provided by the Hive-bee, however much she differs from the mass of Hymenoptera owing to her social habits, her sterile workers and especially her tremendous fertility, extending over so long a period.
I at first doubted the universal presence of this spermatic receptacle, having failed to find it under my scalpel in my former investigations into the anatomy of the Sphex-wasps and some other game-hunters. But this organ is so delicate and so small that it very easily escapes the eye, especially when our attention is not specially directed in search of it; and, even when we are looking for it and it only, we do not always succeed in discovering it. We have to find a globule attaining in many cases hardly as much as a millimetre (12) in diameter, a globule headed amidst a tangle of air-ducts and fatty patches, of which it shares the colour, a dull white. Then again, the merest slip of the forceps is enough to destroy it. My first investigations, therefore, which concerned the reproductive apparatus as a whole, might very well have allowed it to pass unperceived.
In order to know the rights of the matter once and for all, as the anatomical treatises taught me nothing, I once more fixed my microscope on its stand and rearranged my old dissecting-tank, an ordinary tumbler with a cork disk covered with black satin. This time, not without a certain strain on my eyes, which are already growing tired, I succeeded in finding the said organ in the Bembex-wasps, the Halicti (13), the Carpenter-bees, the Bumble-bees, the Andrenae (14) and the Megachiles (15). I failed in the case of the Osmiae, the Chalicodomae and the Anthophorae. Is the organ really absent? Or was there want of skill on my part? I lean towards want of skill and admit that all the game-hunting and honey-gathering Hymenoptera possess a seminal receptacle, which can be recognized by its contents, a quantity of spiral spermatozoids whirling and twisting on the slide of the microscope.
This organ once accepted, the German theory becomes applicable to all the Bees and all the Wasps. When copulating, the female receives the seminal fluid and holds it stored in her receptacle. From that moment, the two procreating elements are present in the mother at one and the same time: the female element, the ovule; and the male element, the spermatozoid. At the egg-layer's will, the receptacle bestows a tiny drop of its contents upon the matured ovule, when it reaches the oviduct, and you have a female egg; or else it withholds its spermatozoids and you have an egg that remains male, as it was at first. I readily admit it: the theory is very simple, lucid and seductive. But is it correct? That is another question.
One might begin by reproaching it with making a singular exception to one of the most general rules. Which of us, casting his eyes over the whole zoological progression, would dare to assert that the egg is originally male and that it becomes female by fertilization? Do not the two sexes both call for the assistance of the fertilizing element? If there be one undoubted truth, it is certainly that. We are, it is true, told very curious things about the Hive-bee. I will not discuss them: this Bee stands too far outside the ordinary limits; and then the facts asserted are far from being accepted by everybody. But the non-social Bees and the predatory insects have nothing special about their laying. Then why should they escape the common rule, which requires that every living creature, male as well as female, should come from a fertilized ovule? In its most solemn act, that of procreation, life is one and uniform; what it does here it does there and there and everywhere. What! The sporule of a scrap of moss requires an antherozoid before it is fit to germinate; and the ovule of a Scolia, that proud huntress, can dispense with the equivalent in order to hatch and produce a male? These new-fangled theories seem to me to have very little value.
One might also bring forward the case of the Three-pronged Osmia, who distributes the two sexes without any order in the hollow of her reed. What singular whim is the mother obeying when, without decisive motive, she opens her seminal phial at haphazard to anoint a female egg, or else keeps it closed, also at haphazard, to allow a male egg to pass unfertilized? I could imagine impregnation being given or withheld for periods of some duration; but I cannot understand impregnation and non-impregnation following upon each other anyhow, in any sort of order, or rather with no order it all. The mother has just fertilized an egg. Why should she refuse to fertilize the next, when neither the provisions nor the lodgings differ in the smallest respect from the previous provisions and lodgings? These capricious alternations, so unreasonable and so exceedingly erratic, are scarcely appropriate to an act of such importance.
But I promised not to argue and I find myself arguing. My reasoning is too fine for dull wits. I will pass on and come to the brutal fact, the real sledge-hammer blow.
Towards the end of the Bee's operations, in the first week of June, the last acts of the Three-horned Osmia become so exceptionally interesting that I made her the object of redoubled observation. The swarm at this time is greatly reduced in numbers. I have still some thirty laggards, who continue very busy, though their work is in vain. I see some very conscientiously stopping up the entrance to a tube or a Snail-shell in which they have laid nothing at all. Others are closing the home after only building a few partitions, or even mere attempts at partitions. Some are placing at the back of a new gallery a pinch of pollen which will benefit nobody and then shutting up the house with an earthen stopper as thick, as carefully made as though the safety of a family depended on it. Born a worker, the Osmia must die working. When her ovaries are exhausted, she spends the remainder of her strength on useless works: partitions, plugs, pollen-heaps, all destined to be left unemployed. The little animal machine cannot bring itself to be inactive even when there is nothing more to be done. It goes on working so that its last vibrations of energy may be used up in fruitless labour. I commend these aberrations to the staunch supporters of reasoning-powers in the animal.
Before coming to these useless tasks, my laggards have laid their last eggs, of which I know the exact cells, the exact dates. These eggs, as far as the microscopes can tell, differ in no respect from the others, the older ones. They have the same dimensions, the same shape, the same glossiness, the same look of freshness. Nor are their provisions in any way peculiar, being very well suited to the males, who conclude the laying. And yet these last eggs do not hatch: they wrinkle, fade and wither on the pile of food. In one case, I count three or four sterile eggs among the last lot laid; in another, I find two or only one. Elsewhere in the swarm, fertile eggs have been laid right up to the end.
Those sterile eggs, stricken with death at the moment of their birth, are too numerous to be ignored. Why do they not hatch like the other eggs, which outwardly they resemble in every respect? They have received the same attention from the mother and the same portion of food. The searching microscope shows me nothing in them to explain the fatal ending.
To the unprejudiced mind, the answer is obvious. Those eggs do not hatch because they have not been fertilized. Any animal or vegetable egg that had not received the life-giving impregnation would perish in the same way. No other answer is possible. It is no use talking of the distant period of the laying: eggs of the same period laid by other mothers, eggs of the same date and likewise the final ones of a laying, are perfectly fertile. Once more, they do not hatch because they were not fertilized.
And why were they not fertilized? Because the seminal receptacle, so tiny, so difficult to see that it sometimes escaped me despite all my scrutiny, had exhausted its contents. The mothers in whom this receptacle retained a remnant of sperm to the end had their last eggs as fertile as the first; the others, whose seminal reservoir was exhausted too soon, had their last-born stricken with death. All this seems to me as clear as daylight.
If the unfertilized eggs perish without hatching, those which hatch and produce males are therefore fertilized; and the German theory falls to the ground.
Mason Bee cocoon
Then what explanation shall I give of the wonderful facts which I have set forth? Why, none, absolutely none. I do not explain facts, I relate them. Growing daily more sceptical of the interpretations suggested to me and more hesitating as to those which I may have to suggest myself, the more I observe and experiment, the more clearly I see rising out of the black mists of possibility an enormous note of interrogation.
Dear insects, my study of you has sustained me and continues to sustain me in my heaviest trials. I must take leave of you for to-day. The ranks are thinning around me and the long hopes have fled. Shall I be able to speak to you again (16)?
1. Between .312 to .468 inch.
2. .195 to .234 inch.
3. 3.9 inches.
4. .195 to .234 inch.
5. .195 to .214 inch.
6. .234 inch.
7. About two-fifths of an inch.
8. .585 inch.
9. .7 inch.
10. .936 inch.
11. Johann Dzierzon, author of „Theorie und Praxis des neuen Bienenfreundes.“
12. About one-fiftieth of an inch.
13. Cf. Chapters 12 to 14 of the present volume.
14. A species of Burrowing Bees.
15. Or Leaf-cutting Bees. Cf. Chapter 8 of the present volume.
16. This is the closing paragraph of Volume 3 of the „Souvenirs entomologiques,“ of which the author has lived to publish seven more volumes, containing over 2,500 pages and nearly 850,000 words.