Bedwyn
Bee.
rish (1851) 5059; of town, 3012, chiefly employed in the manufacture of ribbons, of bricks, and malt. In the vicinity are extensive coal-pits and stone-quarries.
BEDWYN, GREAT, a borough, market-town, and parish in the county of Wilts, 70 miles from London, on the Ken-
net and Avon Canal. It is a place of great antiquity, and, till the Reform bill passed, returned two members to parliament. It has little trade; and the only building that merits attention is the church, which contains several very ancient monuments. Pop. of parish in 1851, 2193.
THE bee, from its singular instincts, its active industry, and the useful products resulting from its labours, has, from the remotest times, attracted the attention, not only of naturalists, but of mankind in general. No nation upon earth has had so many historians as this remarkable tribe of insects. The patience and sagacity of the naturalist have had an ample field for exercise in the study of their structure, physiology, and domestic economy. Their preservation and increase have been objects of assiduous care to the agriculturist; and their reputed perfection of policy and government have long been the theme of admiration, and have afforded copious materials for argument and allusion to the poet and the moralist in every age. It is a subject that has been celebrated and adorned by the muse of Virgil, as well as illustrated by the philosophic genius of Aristotle. Cicero and Pliny report that Aristomachus devoted himself during sixty years to the study of these insects; and Philiscus is said to have retired into a desert wood, that he might pursue his observations on them without interruption. A prodigious number of authors have written express treatises on bees; periodical works have been published relating exclusively to their management and economy; and learned societies have been established for the sole purpose of conducting researches on this subject. The most celebrated association of this kind is the Société des Abeilles, founded about fifty years ago in Little Bautzen, a village in Upper Lausatia, under the auspices of the elector of Saxony. Its labours, as we shall presently find, have enriched the science with a number of valuable discoveries.
In so complicated a branch of natural history, the application of the difficult art of observing correctly, and of the cautious processes of induction, cannot be effected without laborious and long-continued efforts. But, on the subject of bees, the inquirer after truth had, besides, many obstacles to encounter, from the very general diffusion of errors, which had been transmitted without due examination from one author to another. The history of the opinions of successive writers will sufficiently prove how gradual and how slow has been the advancement of real knowledge in what concerns these insects, and will teach us to estimate the value of that which we at length possess, as being the result of the labour of ages, and as being extorted from nature by indefatigable and persevering exertions. So great an accumulation of curious and interesting facts, indeed, has accrued to us from the researches of Swammerdam, Maraldi, Reaumur, Schirach, and Huber, as to constitute almost a new science. Many of these have been discovered subsequently to the time of the compilation of the article BEE in the preceding edition of this work. It will therefore be proper, in this place, to give a connected and systematic account of the natural history of this remarkable insect. For the details of the external characters and distinctions of species, we shall refer to the article ENTOMOLOGY. The principal features of their internal conformation will be described when treating of the particular functions to which they are more immediately subservient; and our descriptions will apply, more especially, to the common and best-known species, the Apis mellifica, which is the one particularly prized on account of the rich products it affords.
The economy of bees comprehends so wide a field of inquiry, the different parts of which are so connected and dependent upon one another, that it is impossible to treat of them distinctly, without supposing the reader to possess some general acquaintance with the history of these insects. We shall therefore premise a brief account of the different sorts of bees inhabiting the hive, and of the respective offices of each. We shall then proceed to consider their comparative physiology; under which head we shall state the leading particulars relating to their nutrition, secretion, respiration, progressive motion, external senses, and instincts. We shall next follow them in their different labours, from the period when the swarm has settled in a new habitation; we shall detail the complex structure of their hives,—their curious processes of architecture,—the pains they bestow on rearing their progeny, and in sending forth new swarms; and this will lead us to the subject of the fecundation of the queens, and the massacre of the drones. After having thus given an account of their usual condition, we shall, in the last place, describe the result of several experiments and observations which have been made when they were placed in unusual circumstances; experiments which have exhibited many interesting features of their character, and have thrown considerable light on the whole of their history, as well as suggested various practical and economic applications in the management of these insects.
The leading feature in their history, and one which distinguishes them from almost all insects, is their singular distribution into three different kinds, constituting, to all appearance, so many different modifications of sex. The drone, which is characterized by a thicker body, a round head, a more flattened shape, and more obtusely-terminated abdomen, within which are contained the male organs of generation, has been admitted as the male of the species. It is distinguished also by the absence of a sting, and by the humming noise that accompanies its flight. The queen-bee, which is larger than any of the others, has the abdomen of greater length, and is provided with a sting, and with two ovaria of considerable size, is unequivocally recognised as the female. The working bees compose the third class, and are distinguished by the smallness of their size, their lengthened proboscis, the peculiar structure of their legs and thighs, which are adapted to the collection of certain materials obtained from vegetables, and by the apparent absence of every trace of generative organs,—we say apparent absence, because, as will be hereafter stated, rudiments of ovaria have been very lately discovered to exist, which, however, are not perceptible without a very minute and careful dissection. Till within a few years the working bees were regarded as animals deprived of sex, and were accordingly termed neuters or mules. It is these which perform all the laborious offices for the community,—which construct the interior of their habitation,—which explore the country in search of nourishment and other materials,—which collect and bring them to the hive, and apply them to different purposes; it is this kind that assiduously attend upon the queen, and supply all her wants,—that defend the hive from the attacks of depredators, and carry on hostilities against the various enemies of the tribe. The life of the
Bee. females is chiefly engrossed with the duties of laying eggs, and conducting the colonies, which, at certain periods, emigrate from the parent state. The drones, producing neither wax nor honey, and depending on the rest for their subsistence, are idle spectators of these labours. They appear to be formed only for the momentary but important duty of impregnation, since they perish when this purpose is accomplished. There is commonly only one perfect queen existing at a time within each hive; and she appears to be treated by all the other bees with every mark of affection and of deference. The number of labourers is very different in different hives; sometimes there are only a few thousands; at other times from twenty to forty, or even fifty thousand. The drones, even in the spring, seldom compose more than one thirtieth or one fortieth of the whole; and, at other seasons, there are none to be found in the hive. In order to form some estimate of the number of bees which can occupy a certain space, Mr Hunter counted what number of drowned bees could be contained in an alehouse pint, and found it to be 2160; so that if a swarm were to fill two quarts, their numbers would be nearly 9000. Reaumur, with the same view of ascertaining their numbers, employed the more accurate method of weighing them; he found that a collection of them, weighing one ounce, consisted of 336 bees; and, therefore, that 16 ounces, or one pound, would consist of 5376 bees.
Real sex of the working bee. Notwithstanding these differences in conformation, instincts, and offices, between the queen-bee and the workers, it is now established, upon the most incontrovertible evidence, that they both originally proceeded from the same kind of larva, and that the queen-bee lays only two kinds of eggs, the one destined to produce the drone, the others capable of being converted, according to circumstances, either into a worker or a queen. It has been proved that the former, although exhibiting no appearance of sexual organs on a superficial examination, are in reality females, and have the rudiments of these organs, which, from their not being developed, are incapable of exercising their proper functions. It may be remarked, that the idea of the working bees being radically females, had been suggested long ago by Dr Warder, in his Monarchy of Bees, in which he terms them "True Amazons;" but no attention had been paid to his opinion. The real merit of this great discovery, which affords the key to a multitude of hitherto inexplicable facts, unquestionably belongs to Mr Schirach, vicar of Little Bautzen, the secretary of the Lausatian Society, to which we formerly adverted. When first announced to the world, it was received with suspicion by the greater number of naturalists, and with complete incredulity by others. It was, indeed, at variance with the whole tenor of the observations of Swammerdam, Maraldi, and Reaumur. Wilhelmi, the brother-in-law of Schirach, though an eye-witness of the experiments from which he had deduced this theory, for a long time refused to admit the doctrine, but became at length one of its most strenuous supporters. It is noticed in a vein of sarcastic ridicule by Mr John Hunter, in his otherwise excellent paper on bees in the Philosophical Transactions. Needham wrote a memoir for the Imperial Academy of Brussels in 1777, for the express purpose of refuting it; and he then inveighs in strong language against those naturalists who had deigned to give it the least countenance. Mr Key, in the Bath Society Papers, declares that he made experiments on this subject for eight years, without obtaining a single result in conformity to Schirach's views. Boonet, after exercising a laudable scepticism, and making a diligent inquiry, in which he displays a genuine spirit of philosophy, yielded a reluctant assent. But the truth of the doctrine has since been placed beyond the reach of
controversy by multiplied series of observations and experiments in different parts of Europe; and more especially by the recent investigations of Mr Huber of Geneva. We shall not at present enter into the detail of proofs, because their force will be better appreciated when other particulars belonging to the history of the bee have been explained.
In considering the physiology of the bee, the first function that claims our notice is that of nutrition. The food of bees is principally of two kinds, namely, the fluid secretions of vegetables contained in the nectarea of the flowers, and the dust of the antheræ, which has been termed by botanists the pollen, but which, when collected by the bees, has received a variety of appellations, such as farina, bee-bread, raw wax (cire brute), &c. Occasionally, however, we find bees feeding upon other saccharine substances besides honey, such as honey-dew, treacle, syrup, &c.
The organs by which they collect food are extremely complex, for they comprise instruments adapted to the reception of liquid aliment as well as those fitted for the division of solid materials. Reaumur has given the most elaborate description of these organs, in which he has corrected some errors that Swammerdam had fallen into. For the purpose of taking up fluids, they are provided, in common with all hymenopterous insects, with a long and flexible proboscis or trunk, which may be considered as a Proboscis. lengthened tongue, though, strictly speaking, it is formed by a prolongation of the under lip. It is not tubular, as Swammerdam had supposed, but solid throughout; and the minute depression at its extremity is not the aperture of any canal through which liquids can be absorbed. Cuvier, in his Leçons d'Anatomie Comparée, has not marked this distinguishing feature in the proboscis of the bee, but speaks of it in common with the tubular trunks of the other hymenoptera, and describes its aperture as being situated in the lower part. But Reaumur has very satisfactorily shown that the trunk of the bee performs strictly the office of a tongue, and not that of a tube for suction; for when it takes up honey or other fluid aliment, the under or the upper surfaces are more immediately applied to it, and rolled from side to side, and the bee thus licks up what adheres to it, while the extremity of the trunk is frequently not applied at all to the substance taken up. The trunk is supported on a pedicle, which admits of being bent back or propelled forwards, and thus can retract or stretch out the trunk to a considerable extent. Protection is given to it by a double sheath; the external, consisting of two scales furnished by the expansion of one of the portions of the labial palpi; and the internal, formed by the prolongation of the two external portions of the jaw. The whole member thus consists of five principal parts, on which account Fabricius termed it lingua quinquefida.
For the purpose of mechanically dividing solid mate- Mandibles. rials, the mouth is furnished with two strong mandibles and four palpi; they are but little employed in eating, but are of great use in enabling the insect to seize and break down hard substances for other purposes. In the working bee all these parts are of larger dimensions than in the other kinds. The teeth are two in number, and have the form of concave scales with sharp edges; they are fixed to the ends of the jaws, and play horizontally as in other insects. Reaumur describes and delineates a large aperture above the root of the proboscis, which is so surrounded with fleshy parts as not to be readily seen unless the proboscis be extended and bent downwards. This he considers as the mouth or orifice of the gullet; on the upper side of which, and of course opposite to the root of the proboscis, a small fleshy and pointed organ is seen, which he regards as the tongue, assisting in the deglutition of the
food. Through this orifice, it is presumed, all the aliment, whether liquid or solid, passes; the former being conveyed to it by the trunk, which, by its contractile power, presses forward the fluids it has collected between itself and the inner sheath, and the latter being received directly after its comminution by the teeth, behind which it is situated. Latreille, however, whose authority is great on a point of this nature, thinks that Reaumur has deceived himself with regard to such an aperture, and disbelieves its existence. He conceives that the food simply passes on by the sides of the tongue, finding its way from thence into the œsophagus, and so on to the stomach.
The bee has two stomachs; the first is a large transparent membranous bag, pointed in front, and swelling out into two pouches behind. It performs an office in some respects analogous to that of the crop in birds; for it receives and retains for a time the fluid of the nectarea, which does not appear to differ in any respect from honey. Mr Hunter observes, that, whatever time the contents of this reservoir may be retained, he never found them altered, so as to give the idea of digestion having taken place. The coats of this reservoir are muscular, by which means it is capable of throwing up the honey into the mouth, so that it is regurgitated into the honey cells, or imparted to other bees. None of it ever passes out from the extremity of the trunk, as Swammerdam had believed. For the purpose of digestion, a second stomach is provided, which takes its origin from the middle of the two posterior lobes of the former, and is of a lengthened cylindrical shape. Its communication with the intestine is not direct, but takes place by a projecting or inverted pylorus, thickest at its most prominent part, with a very small opening in the centre, of a peculiar construction. This inward projecting part is easily seen through the coats of the reservoir, especially if full of honey. A similar kind of structure takes place at the communication of the first with the second stomach, and, having the properties of a valve, must effectually prevent all regurgitation from the latter into the former.
The pollen of flowers, which is the other principal article of food, was shown by Swammerdam to consist of an infinite number of small particles, generally of a globular shape, each of which is found to be a small capsule, enclosing the still finer dust or fecundating principle destined to be shed on the pistils for the purpose of germination. Geoffroy has given a memoir, published in the Collection Académique des Sciences, containing a minute description of the shapes of these capsules, taken from different flowers. The working bees, by means of the pencil of hair which grows on the tarsi, first collect a certain quantity of pollen, which they knead together into a ball, and place it in the concave space which is situated at the middle joint of the hinder feet, and has been termed the basket. The surrounding rows of hairs keep the ball from falling off. In order to gather larger quantities at once, the bees are sometimes observed to roll their bodies on the flower, and then, brushing off the pollen which adheres to them with their feet, form it into two masses, which they dispose of as before mentioned; and it is said that, in moist weather, when the particles of pollen cannot be readily made to cohere together, they return to their hive dusted all over with pollen, which they then brush off with their feet. They are often obliged to tear open the capsules which contain the pollen, in order to procure a supply of this substance, when it has not yet been shed by the flowers.
Pollen is yielded by flowers during the spring in such abundance that the bees of a single hive will often bring back one pound, or even more, in a day. Some agriculturists have accordingly imagined, that the vegetation of
some plants might be endangered from this great consumption of the fecundating principle by insects in general; for other insects besides bees seek it with avidity. But this fear has been proved to be totally without foundation, and the practice of destroying bees in order to prevent this imaginary danger is therefore as useless as it is barbarous. It would appear, indeed, that so far from obstructing the fecundation of plants, the labours of the bee have often tended materially to promote it, by the agitation which they give to the flower, and by transporting the pollen from one flower to another. In this manner may we account for the number of hybrid flowers that are met with near the haunts of bees.
It has been shown very clearly by Huber, in a paper in the Journal de Physique, that pollen is peculiarly the food of the young bees, and is collected by the working bees with this intention. Reaumur, however, asserts that he has seen adult bees devour pollen. Swammerdam, who conceived the trunk to be tubular, rejected the idea that pollen could ever be the food of bees, as the globules of which it consists are incapable of entering an orifice so minute as that which appears at the extremity of the trunk, and which, as he was unacquainted with the real mouth, he thought was the only passage to the stomach. Latreille, who does not admit the existence of the large mouth described by Reaumur, states that the mandibles lay hold of the pollen, and carry it to the base of the trunk, from whence it finds its way into the œsophagus by the sides of that organ.
Of honey-dew, which forms part of the food of bees, we shall merely remark that it is an excrementitious matter deposited on the leaves of plants by certain species of aphides.
An abundant supply of water is essential to the healthy condition of bees. They consume a large quantity, and often stop to drink at the edge of stagnant pools, and seem even to prefer putrid and urinous waters to purer streams, as if their saline and pungent qualities were grateful to them.
It has been long the opinion that wax was but a slight modification of pollen, which required for this conversion of wax merely the application of a certain pressure, and a kind of kneading by the feet of the bees. Many naturalists, such as Bernard de Jussieu, had persuaded themselves that the dust of the stamens of flowers contained wax ready formed as one of its ingredients, and quoted the following experiment in proof of this opinion: If the minute grains of pollen be put into water, they gradually swell, till they at length burst, at which moment a small jet of an oily liquor will be perceived, which floats on the water without mixing with it. But Reaumur had attempted in vain to extract any thing like wax from dust of the anthers; and, indeed, an attention to the chemical properties of these two substances would have sufficiently pointed out their essential differences. From the upper surface of the leaves of many kinds of trees, a substance has indeed been obtained, which possesses all the qualities of bees' wax; but nothing like it can be extracted from pollen. Reaumur was persuaded that the pollen was elaborated in the second stomach of the bee, and thrown up into the mouth in the form of a white foam, which, by exposure to the air, hardened, and became wax; and that the bee took advantage of its soft state to apply it in the building of the combs. So circumstantial an account, given to us by a scrupulous observer of facts, appeared to be perfectly satisfactory, and was acquiesced in by naturalists in general. But it has since been completely proved by the researches of Duchet, of Hunter, and of Huber, but principally by the latter, that wax is a secretion from the abdomen of the bee; and that it depends not at all
Bee. on the pollen which the insect may consume, but on the quantity of honey or other saccharine substance which it receives into the stomach. The first step in this discovery was made by one of the members of the Lausian Society, whose name has not been preserved. It was mentioned in a letter of Mr Wilhelmi to Bonnet, in August 1768, in which he says that wax, instead of being rejected by the mouth, exudes from the rings which enclose the posterior part of the body. Of this we may satisfy ourselves by drawing out the bee from the cell in which it is working with wax, by means of the point of a fine needle; and we may perceive, in proportion as the body is elongated, that the wax will make its appearance under the rings, in the form of small scales. Mr Duchet, in his Culture des Abeilles, gives a full statement of the principal circumstances attending the production of wax, which he very justly ascribes to the conversion of honey into this substance in the body of the bee. These facts appear to have been entirely overlooked till the subject was again brought forward by Mr John Hunter, in his paper in the Philosophical Transactions for 1792. Wildman, however, had cursorily remarked that portions of wax, in the form of scales, and which he conceived must have been moulded on the body of the bee, are sometimes found at the bottom of the hive. Mr Huber was engaged in prosecuting his inquiries on this subject at the same period with Mr Hunter, and discovered, in 1793, the existence of regular receptacles or pouches, from the coats of which the wax is secreted, and within which it accumulates till its edges raise the scales, and become apparent externally. These plates of wax are withdrawn by the bee itself, or some of its fellow-labourers, and are applied in a manner hereafter to be described.
Huber has shown, by a series of well-conducted experiments, that, in a natural state, the quantity of wax secreted is in proportion to the consumption of honey; but that an equal or even greater quantity will be formed if the bee be fed on a solution of sugar in water. Warmth and rest promote this process of secretion; for the bees, after feeding plentifully on saccharine food, hang together in a cluster without moving, for several hours, at the end of which time large plates of wax are found under the abdominal rings. This happened when bees were confined and restricted from any other sort of nourishment; whilst those that were fed on pollen and fruits alone did not produce any wax. It appears also from his researches that the formation of wax is the office of a particular set of bees, which may be distinguished from the rest, and particularly from those that nurse the young larvae, by the greater size and more cylindrical shape of their abdomen. Dissection also shows that their stomachs are more capacious. In the second volume of Huber's Nouvelles Observations sur les Abeilles, he describes minutely the anatomy of the pouches or receptacles for the wax, which are parts peculiar to the working bees, being totally absent in the males and queens. It is a structure that had escaped the keen eyes of Swammerdam, and has not been noticed by any subsequent anatomist. The cavities are lined with a membrane, which presents a number of folds, forming an hexagonal net-work, not unlike the appearance in the second stomach of ruminant quadrupeds, and evidently destined to perform the office of secretion.
Among the secretions peculiar to the bee, the poison which is poured into the wounds made by the sting deserves to be noticed. It is said to owe its mischievous efficacy to certain pungent salts. If a bee is provoked to strike its sting against a plate of glass, a drop of poison will be discharged; and if this is placed under a microscope, the salts may be seen to concretize, as the liquor dries, into clear, oblong, pointed crystals. The sting con-
sists of bearded darts, which are protruded from the end of a sheath; and the venomous juice is likewise injected through the sheath, from a little bag at the root of the sting.
As no organs for the circulation of blood have been as Function yet demonstrated in insects, respiration is supposed to be of respiration effected by means totally different from those which are adopted in the higher classes of the animal kingdom. As the blood, or fluid corresponding to the blood, cannot be presented to the air in any separate organ, the air must be conducted to the blood, wherever such a fluid is met with. For this purpose, tracheæ or air-tubes, having several external openings or spiracles, are made to ramify like arteries, and are distributed in an infinite number of branches to every part of the body. The analogy of other insects might perhaps be admitted as sufficient evidence that bees respire atmospheric air, the constant renewal of which is essentially necessary to the continuance of the vital functions. It is, however, not always safe to trust to analogical reasoning in subjects of natural history; and direct evidence is, in all cases, to be preferred when it can be obtained. We must therefore consider as valuable the complete series of experiments on the respiration of bees, that have been lately given to the world by Huber, to whom we already owe so large a portion of the information we possess with regard to these insects. We might indeed have anticipated, with the strongest probability, many of the results to which these experiments have led; but there are others which are quite unexpected, and possess as much of interest as of novelty.
The condition of a hive of bees, in which many thousand individuals, full of animation and activity, are crowded together in the very small space of one or two cubic feet, having no communication with the external air but by means of a small aperture in the lowest part, which entrance is frequently obstructed by a throng of bees that are passing in and out during sultry weather, is of all possible conditions the one least favourable to the renewal of heated air. The most crowded theatres or hospitals are not to be compared with it in point of closeness. Direct experiment, indeed, shows that the combustion of a taper could not be carried on in so limited a space; for Mr Huber found that, in a glass ball of the same dimensions as the hive, and with a similar aperture, the taper went out in a few minutes. So great was the difficulty of explaining the respiration of bees under these circumstances, that Mr Huber was led to examine into the truth of the opinion, that respiration was equally necessary to its bees as to other insects. The results were unequivocal. They perish speedily in the vacuum of the air-pump. They are easily drowned by placing them so that the spiracles on the corslet are under water; but revive readily when they are dried. The action of the spiracles is, in this experiment, rendered manifest by the escape of bubbles of air from each of their orifices. When a number of bees are confined in a bottle accurately closed, they exhibit unequivocal symptoms of distress, and fall into complete asphyxia. These changes occur more rapidly when they are placed in any gas which contains no admixture of oxygen, such as carbonic acid, hydrogenous and azotic gases. When they are rendered torpid by cold, and respiration is thereby suspended, these effects do not take place. All these effects are more considerable in adult bees than in the larvae, though they are also distinctly exhibited in the latter. Suffocation is retarded if the proportion of oxygen be greater than in atmospheric air, and it may be averted altogether by a continual renewal of oxygen. It was ascertained by the eudiometer that the same changes were produced on the air as in the respiration of other animals; namely, the subtraction of
Bee. oxygen and the addition of an equal volume of carbonic acid gas.
Yet, on examining the air of the hive itself, it was found scarcely to differ in purity from atmospheric air. It was at one time conjectured that some of the contents of the hive, such as the pollen, the honey, or the wax, might have some power of evolving oxygen, so as to afford the requisite supply of this gas. Experiments, however, proved that they had no such power. Amidst so great an uncertainty, it was thought worth while to ascertain whether bees might not exert some unknown process by which oxygen was generated in the hive itself. If this were true, they could support life although all communication with the external air were intercepted. A hive was selected having glass sides, so as to allow of the observer's seeing what was passing in the interior, and the entrance was completely closed. In a quarter of an hour the bees became sensible of their situation, and showed great uneasiness; all business was suspended; an extraordinary agitation, accompanied by a remarkable noise, prevailed in every quarter. All the bees were seen beating their wings with the same rapidity as in flying. They were thus incessantly occupied during ten minutes. Their motions became then more languid, and, after being utterly exhausted, they fell in succession to the bottom of the hive, till every one of them was in a state of complete asphyxia. It is remarkable that, at this period, the temperature of the hive, which had been previously at 95° of Fahrenheit, suddenly cooled down to that of the external air. On opening the door and the top of the hive, and establishing a current of air through it, the bees were soon restored to animation.
It was proved by this experiment that the air is renewed through the small opening which serves as a door to the hive. By suspending light substances near the entrance, the existence of different currents of air was rendered manifest. After much reflection, it occurred that the violent agitation of the wings might have some influence in procuring this renewal of air. This conjecture was confirmed by an experiment with a glass bell, to the aperture of which an apparatus was fitted, consisting of a small ventilator, which could be moved rapidly round by machinery. When the ventilator was set in motion, the air within could support the combustion of a candle for an unlimited time. Observation further showed that some bees are actually always employed in the office of ventilating the hive; they vibrate their wings with great vigour and constancy, producing so rapid a movement of them, that they cannot be seen except in the two extremities of the arc of vibration, which is at least one of 90°. While thus imitating the actions of flying, they fasten themselves with their feet to the floor of the hive, so that the whole effect of that impulse which, were they at liberty, would carry them forwards with considerable velocity, is exerted on the air, which is therefore driven backwards in a powerful current. Some bees occasionally perform these ventilating motions on the outside of the hive, near the entrance; but a still greater number are employed in this office within doors: sometimes twenty are thus occupied at once, and each bee continues its motions for a certain time, occasionally for nearly half an hour, and is then relieved by another, who takes its place. This is the occasion of that humming sound which is constantly heard from the interior of the hive when the bees are not in a state of torpidity. But it is often heard with even more than usual loudness in the depth of winter. The warmth of the sun's rays, however, always occasions an increased activity among the ventilating bees. The immediate cause of these actions is probably some impression made on their organs by the presence of vitiated air; for a bee
may be made to ventilate itself by placing near it substances which have to them an unpleasant odour, such as spirit of wine or oil of turpentine.
The connection between an active respiration and a high temperature is remarkably exemplified in bees, among which, in consequence of their collecting together in large numbers, the heat is not so easily dissipated, and admits also of being easily ascertained by the thermometer. Mr Hunter found it to vary from 73° to 84° of Fahrenheit; and Mr Huber observed it on some occasions to rise suddenly from about 92° to above 104°.
Bees are well fitted, by their structure, for rapid flight through the air. They possess great muscular strength in proportion to their size, and their indefatigable activity in the different labours of the hive is truly astonishing. Aristotle and Pliny have pretended that, during high winds, they endeavour to steady their flight by holding a small stone with their feet, by way of ballast. This assertion has been shown, both by Swammerdam and Reaumur, to be erroneous in as far as it applied to the common bee; but there are other species which build nests with stones and other hard materials, and which, while transporting them for this purpose, were probably mistaken for the honey-bee.
The physiology of the external senses must necessarily be very imperfectly understood in a class of animals of a nature so remote from our own species. The infinite diversity of characters presented to us by the different tribes of insects, as well as of other animals, naturally suggests the idea that external objects produce on their sentient organs impressions widely different from what they communicate to ourselves. The notions we form of their senses must not only be liable to great inaccuracy, but must often be totally inadequate representations of the truth. A finer organization, and more subtle perceptions, would alone suffice to extend the sphere of their ordinary senses to an inconceivable degree, as the telescope and the microscope have with us extended the powers of vision. But they possess, in all probability, other organs, appropriated to unknown kinds of impressions, and which must open to them avenues to knowledge of various kinds, to which we must ever remain total strangers. Art has with us supplied many elaborate modes of bringing within our cognizance some of the properties of matter which nature has not immediately furnished us with the means of detecting. But who will compare our thermometers, electroscoops, or hygrometers, however elaborately constructed, with those refined instruments with which the lower classes, and particularly insects, appear to be so liberally provided? The antennæ, which are so universally met with in this class of animals, are doubtless organs of the greatest importance in conveying impressions from without. Their continual motion, the constant use which is made of them in examining objects, the total derangement in the instincts of those insects which have been deprived of them, point them out as exquisite organs of more than one sense. To impressions of touch, arising from the immediate contact of bodies, they are highly sensible; but their motions evidently show that they are affected by objects at some distance. They are no doubt alive to all the tremulous motions of the surrounding air, and probably communicate perceptions of some of its other qualities. Composed of a great number of articulations, they are exceedingly flexible in every direction, and can readily embrace the outline of any body that the bee wishes to examine, however small its diameter, and are capable of following all its movements. It is by means of these instruments that the bee is enabled to execute so many works in the interior of the hive, from which the light must be totally excluded.
Bee. ed. Aided by these, it builds its combs, pours honey into its magazines, feeds the larvæ, and ministers to every want, which it discovers and judges of solely by this species of touch.
The antennæ appear also to be the principal means employed for mutual communication of impressions. The different modes of contact constitute a sort of language, which appears to be susceptible of a great variety of modifications, and to be capable of supplying at once every species of information for which they have occasion. It is in this way alone that they satisfy themselves of the presence of their queen, or communicate to others the alarming intelligence that she has disappeared.
Vision. The sense residing in the antennæ appears to be, on many occasions, supplementary to that of vision, which in bees, as in other insects, is less perfect than in the larger animals. During the night, therefore, they are chiefly guided in their movements by the former of these senses. This will sufficiently appear from observing by moon-light the mode in which the bees guard the entrance of the hive against the intrusion of moths which flutter in the neighbourhood. They act as vigilant sentinels, performing continual rounds near this important post, extending their antennæ to the utmost, and moving them alternately to the right and to the left. Woe to the unfortunate moth that comes within their reach. Aware of its danger, and of the defective sight of the bees, the moths adroitly avoid the slightest contact, and endeavour to insinuate themselves between the bees, so as to get unperceived into the hive, where they riot upon the honey which they find.
If bees require full day-light for the exercise of vision, it must at the same time be acknowledged, that, when they are so assisted, they appear to enjoy this sense in great perfection. A bee will recognise its habitation from great distances, and distinguish it at once from many others in a numerous apiary. It passes through the air in a straight line towards its object with extreme rapidity. On quitting the hive, it flies towards the field which is most in flower; and as soon as it has determined on its course, it takes as direct a line as a ball issuing from a musket. When it has collected sufficient provision, it rises in the air to discover its hive, and then darts forward with the velocity of an arrow, and with unerring precision in its aim.
Perceptions of temperature. Their perceptions of heat and cold, which are generally referred to the sense of touch, appear to be extremely delicate. In several experiments of Huber's, the influence of the rays of the sun excited them to a vigorous action of the wings. It is well known that great cold reduces them to a state of torpor, and inferior degrees of cold are evidently unpleasant to them. They show by their conduct that they are sensible of alterations in the state of the weather for some time before we can perceive them. Sometimes, when working with great assiduity, they will suddenly desist from their labours; none will stir out of the hive, while all the working-bees that are abroad hurry home in crowds, and press forward so as to obstruct the entrance of the hive. Often when they are thus warned of the approach of bad weather, we can distinguish no alteration in the state of the atmosphere. Gathering clouds sometimes produce this effect on them; but perhaps they possess some species of hygrometrical sense, unconnected with any impression of vision. It is alleged that no bee is ever caught in a sudden shower, unless from some cause it has wandered very far from the hive, or been disabled by some accident from returning to it. There is reason for thinking, however, that much exaggeration has prevailed in the statements of authors as to the extent of this kind of foresight. Huber supposes that it is the rapid diminution of light that alarms them; for if the sky be uniformly overcast, they proceed on their excursions, and even the
first drops of a soft shower do not make them return with any great precipitation.
Taste. Their taste is perhaps the most imperfect of their senses. Taste. They exert hardly any discrimination in the collection of honey from different flowers. They are not repelled by the scent or flavour of such as are extremely offensive to our organs, and scruple not to derive supplies from such as are highly poisonous. In some districts in America, it is well known that the honey acquires in this way very deleterious properties. The qualities of honey are, indeed, observed to vary much, according to the particular situation from which it is obtained. The most stagnant and putrid waters, as we have already noticed, are resorted to by bees with the same avidity as the purest. In their selection of flowers they are guided by the quantity of honey they expect to meet with, and in no respect by its quality. When the scythe has cut down all the flowers which before yielded them a plentiful supply, they discontinue their excursions, although the weather be in all respects propitious. Their smell must, therefore, be sufficiently acute to enable them to discover the presence of honey at great distances. Direct experiment has, indeed, proved this to be the case. Mr Huber found that they proceeded immediately towards boxes which contained honey concealed from their view; and such in fact is the situation of the fluid of the nectaræa in flowers. Some odours, especially the fumes of tobacco, and indeed all kinds of smoke, are highly obnoxious to them; this is the case, also, with the smell of oil of turpentine, alcohol, ammonia, the nitric and muriatic acids, and several other volatile chemical agents, upon receiving the impressions of which, they immediately set about ventilating themselves in the manner above described. But nothing excites their displeasure in a greater degree than the breath of the spectator; as soon as they feel which, they show signs of anger, and prepare to revenge it as an insult. The odour of the poison of their sting produces similar effects, exciting them to immediate rage and hostility.
Smell. Although it is sufficiently clear that many insects possess the power of smell, yet the particular organ of this sense has never been accurately ascertained; and the opinions of naturalists have been much divided on the subject. These opinions have been supported more by arguments drawn from the analogy of what happens in other classes of animals, than by any direct experiments on insects themselves. We know that, in all animals respiring by means of lungs, the organs of smell are placed at the entrance of the passages of the air; and it has often been concluded, that in like manner the stigmata, or the orifices of the air-tubes, were the seat of this sense in insects. By others the antennæ have been assigned as the organs through which these impressions were conveyed to the sensorium. The experiments of Huber have proved that neither of these opinions is correct; and have satisfactorily shown that in the bee this sense resides in the mouth itself, or in its immediate vicinity. Here, indeed, would be its proper station, if this faculty be intended, as we may reasonably suppose it to be, to apprise the individual of the qualities of the food prior to its being eaten. When the mouth of the bee was plugged up with paste, which was allowed to dry before the insect was set at liberty, it remained quite insensible to the same odours to which it had before manifested the strongest repugnance.
It is generally supposed that bees possess the sense of Hearing. hearing. The common practice of making a loud noise by drums and kettles in order to attract a swarm is founded on this supposition. But the evidence is by no means conclusive; for we find that they are nowise disturbed by a loud clap of thunder, or by the report of a gun, or any
other noises that may happen to arise around them. It is, however, certain that they are capable of emitting a variety of sounds, which appear expressive of anger, fear, satisfaction, and other passions; and it would seem that they were even capable of communicating certain emotions to one another in this manner. Huber observed that the queens, during their captivity, sent forth a peculiar sound, which he supposes to be a note of lamentation. A certain cry or humming noise from the queen will strike with sudden consternation all the bees in the hive; and they remain for a considerable time motionless and stupified. Hunter has noticed a number of modulations of sound emitted by bees under different circumstances, and has instituted an inquiry concerning the means employed by them in producing these sounds; for an account of which we shall refer the reader to his paper in the Philosophical Transactions.
If the function of sensation in insects be involved in doubt and obscurity, the knowledge of those more interior faculties which are the springs of voluntary action is hid in still deeper mystery. Buffon refuses to allow bees any portion of intelligence, and contends that the actions we behold, however admirably they are directed to certain ends, are in fact merely the results of their peculiar mechanism. Other philosophers, such as Reaumur, have gone into the opposite extreme, and have considered them as endowed with extraordinary wisdom and foresight, as animated by a disinterested patriotism, and as uniting a variety of moral and intellectual qualities of a higher order. The truth, no doubt, lies between these overstrained opinions; but it is nevertheless extremely difficult to decide in what degree these respective principles operate in the production of the effects we witness. We have been too long in the habit of sheltering our ignorance of the causes of this class of phenomena, by referring them indiscriminately to what is called instinct, to submit to a cautious and patient investigation of the hidden springs of action. The term instinct should properly be regarded, not as denoting a particular and definite principle of action, whose operation we can anticipate in any new or untried combination of circumstances, but as expressive of our inability to refer the phenomena we contemplate to any previously known principle. Thus the actions which an animal performs in obedience to the calls of appetite are not properly said to be instinctive; nor can the term be applied to actions which are the consequence of acquired knowledge, and of which the object is with certainty foreseen by the agent. But when an animal acts apparently under a blind impulse, and produces effects useful to itself or to the species, which effects it could not have previously contemplated as resulting from those actions, it is then customary to say that it is under the guidance of instinct, that is, of some unknown principle of action. It will be proper, therefore, to keep this distinction in view in judging of the voluntary actions of the lower animals.
In no department of natural history is it more necessary to be aware of the proper import of the term instinct, than in studying the phenomena presented by the bee; for nowhere is it more difficult to discriminate between the regular operation of implanted motives, and the result of acquired knowledge and habits. The most striking feature of their history, and the one which apparently lays the foundation for those extraordinary qualities which raise them above the level of other insects, is the disposition to social union. It may in general, indeed, be remarked, that animals which associate together so as to form large communities, display a higher degree of sagacity than those which lead a solitary life. This is especially observable among insects. The spider and Formica leonis may exhibit particular talents, or practise particular stra-
tagems in the pursuit and capture of their prey; but their history is limited to a single generation, and embraces none of those interesting relations which obtain between individuals composing the gregarious tribes, such as the ant, the wasp, and the bee. Among these we trace a community of wants and desires, and a mutual intelligence and sympathy, which lead to the constant interchange of good offices, and which, by introducing a systematic division of labour, amidst a unity of design, leads to the execution of public works on a scale of astonishing magnitude. The attachment of bees to their hive, which they defend with a courage and self-devotion truly admirable; their jealousy of intruders; their ready co-operation in all the labours required for the welfare of the community; their tender care of their young; the affection and homage which they bestow on their queen, and which they manifest on all occasions in the most unequivocal manner; imply qualities such as we could hardly persuade ourselves could animate a mere insect, on which we are in the habit of proudly looking down as placed in one of the lowest orders of created beings.
We shall content ourselves at present with these general observations, as the instances which serve to illustrate their moral and intellectual character belong properly to the history of the different processes they follow in the construction of their combs, the hatching and rearing of their progeny, and the mode of conducting their migrations. To these subjects, therefore, we shall now proceed: and in order to present the most connected and complete account of their economy, we shall begin the history from the period when a new swarm has just occupied a hive, and when all the arrangements for their habitation, and the construction of the cells in which their eggs and provisions are to be deposited, are yet to be effected.
The first care of the labouring bees, on their settlement Preparation of the hive. in their new abode, is to clean it out thoroughly. While one set of bees is thus employed, another is distributed about the country in order to procure the proper materials for blocking up the small holes and chinks of the hive, and for laying a firm foundation for the edifice which is to be constructed within it. The substance which is principally employed in this preliminary stage is propolis, a species of glutinous resin, of an agreeable aromatic odour, and reddish-brown colour, in process of time becoming darker, and acquiring a firmer consistence. According to the analysis of Vauquelin (Mém. Soc. Agricult. Departem. Seine), it is composed chiefly of resin, with a small proportion of wax, and of acid and aromatic principles. It is soluble in alcohol, ether, and oils, both fixed and volatile; and tinges the solvent of a beautiful red colour. Cadet has since ascertained in it the presence of benzoic and gallic acids. Reaumur had not been able to discover from what plants the bees collect this substance. Riem asserts that it is chiefly from pines and other trees of the fir kind. The recent observations of Huber have assisted in the solution of this question. On placing branches of the wild poplar tree before the hive, he found that the bees eagerly seized upon the varnish which exudes from the buds; and examining the chemical properties of this varnish, he identified it with the propolis with which the inside of the hive is lined.
The propolis adheres so strongly to the legs and feet of Mode of Mode of the bee which has collected it, that it cannot be detached its application without the assistance of its fellow-labourers. For this purpose the bee that is loaded presents its legs to the workers in the hive, which carry off with their jaws this adhesive substance, and immediately apply it, while yet ductile, all round the interior of the hive, and particularly over all the projecting parts; hence its name, of Greek