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VOL. I.

Bees and Bee-keeping.

SCIENTIFIC AND PRACTICAL.

Plate : /.

Digestive System of Bee (Magnified Ten times).

A, Horizontal Section of Body— Ip, Labial Palpus ; mx, Maxilla; e. Eye; dv, dv,
Dorsal Vessel; v. Ventricles of the same; No. 1, No. 2, No. 3, Salivary Gland
Systems, 1, 2, 3 ; oe, (Esophagus; pro.t, Prothorax ; mesa.t, Mesathorax ; meta.t,
Metathorax; g , g, Ganglia of Chief Nerve Chain; n, Nerves; hs. Honey
Sac; p, Petaloid Stopper of Honey Sac or Stomach-Mouth; c.s, Chyle
Stomach; bt, Biliary or Malpighian Vessels; si, Small Intestine; l, Lamellae
or Gland Plates of Colon ; li, Large Intestine. B, Cellular Layer of Stomach—
gc, Gastric Cells, magnified 200 times. C, Biliary Tube—be, Bile Cells;;,
Trachea. D, Inner Layer, carrying gt, Gastric Teeth.

BEES & BEE-KEEPING;

Scientific anb practical.

A Complete Treatise on the Anatomy, Physiology,
Floral Relations, and Profitable Management
of the Hive Bee.

FRANK R. CHESHIRE, F.L.S., F.R.M.S.,

Lecturer on Apiculture at South Kensington,

Author op “Diagrams on the Anatomy of the Honey Bee;” “Practical
Bee-Keeping;” “Abdominal Distention in Bees during Winter;”
“Honey as Food;” “The Apparatus for Differentiating the
Sexes in Bees and Wasps;” “The Relations of Insects to
Flowering Plants;” “Foul Brood not Micrococcus
but Bacillus—The Means of its Propagation,
and the Method of its Cure;” etc., etc.

With Numerous Illustrations of the Internal and External Structure
of the Bee, and its Application to Plant Fertilisation ;

Bee Appliances , and Methods of Operation, Diseases, &c.,
expressly drawn for this work by the Author.

VOL. I —SCIENTIFIC.

LONDON:

L. UPCOTT GILL, 170, STRAND, W.C.

All Bights Reserved.

PREFACE.

I N deference to repeated requests, made by many
unknown correspondents, as well as by those who
have listened to his Courses of Lectures, the Author
has prepared the following pages for publication ; and,
in presenting them to the attention of the Scientific
and Bee-keeping public generally, he also fulfils a
promise, given ten years since, in his “ Practical

Bee-keeping,” for the very flattering reception of
which he now tenders thanks to his numerous
readers.

The unique character of the present work will
explain and excuse the delay in its appearance;
for the Author has never scrupled to devote time
without stint to the solution of any difficulty,
and has preferred, in every possible case, to put

generally received statements to the test of careful
experiment before adopting them as his own.
This has led to numerous discoveries, which alto¬
gether change the aspect of some parts of the

subject; while many old landmarks have had to

submit to modifications in their positions, or absolute
displacement. The Author acknowledges that he
possesses rather the spirit of the progressionist and
investigator than that of the antiquarian, and, should
he in any case be suspected of having been led

PREFACE.

into a too hasty substitution of a new theory for
an old, he begs no other indulgence, than that his
conclusions should be tested with care at least
equal to that by which he has endeavoured to
support them. He thus trusts that, truth prevailing,
the little corner of the vast field of human inquiry
in which he has had the honour to labour may be
enriched by facts which may give a new departure,
and make Apiculture both more delightful and more
profitable, because more intelligent.

It has been thought wise to treat, as far as
possible, the Scientific and Practical aspects of the
question, notwithstanding the close dependence of the
latter upon the former, in two separate volumes,
which should be, together, a complete guide in all
matters, both touching the Natural History and Man-
agement of the Hive Bee. Of these volumes, the
first is mainly devoted to the consideration of the
anatomy and physiology of the bee itself, the
peculiarities of the sexes, and the principles of
comb structure, while the fascinating botanical ques¬
tion of the relation of bees to flowering plants has
been rather fully treated. The Illustrations here are,
in some points, like those in the Author’s large
Diagrams, published by the British Beekeepers’ Asso¬
ciation, and which, unfortunately, have been copied
by one or two writers with a faithfulness which is
flattering, although, with a reticence far more common
than commendable, they have abstained from men¬
tioning the source whence their material has been

PREFACE.

vii

derived. The second volume deals directly and
very fully with Practical and Commercial matters,
but always in the scientific spirit, for this separation
has been carried so far only as the convenience of
the reader seemed to demand— e.g., chemical and
chemico-physiological matters touching the nature
of honey—its value as a human food; its adultera¬
tion, and tests for the same ; artificial foods for bees,
and their essentials ; the characteristics of pure bees’
wax, and many other such, beside the highly im¬
portant, and now extensive, question of Bee Diseases,
and their Treatment, the latter being necessarily
discussed in the light of the microscopical dis¬
coveries, upon which all knowledge of the subject
really rests.

Investigators, discoverers, and inventors have been
duly credited, and all sources of information stated;
the works also, within the knowledge of the Author,
which might be useful to the reader desiring to
prosecute any special point, have had their titles
given in full in footnotes.

While coveting for this effort the encouragement
which former ones have received, the Author desires
to acknowledge the assistance given by his Pub¬
lisher, who has aided him in every endeavour to
make “ Bee-keeping, Scientific and Practical,” worthy
of support; and to a loving Daughter, who has
read his proofs, it is a deep pleasure to express his
indebtedness.

Avenue House, Acton, W.,

January, 1886 .

CONTENTS.

CHAP. PAGE

Introduction .i

I. —Wild and Hive Bees.7

II. —Economy of Hive Bee.15

III. —General Structure.29

IV. —Nerve System.45

V.—Digestive System.57

VI.—Salivary (?) Glands of Bees.72

VII.—Tongue and Mouth Parts.87

VIII.—Organs of Special Sense : Antennae and Eyes - 102

IX. —Thorax and Legs. 120

X. —Wings and Flight : Buzzing and Humming - - 134

XI. —Secretion of Wax, and Bee Architecture - - 151

XII.—Sting Structure.183

XIII. —Organs of the Drone. 198

XIV. —Queen Organs and Development - - - - 212

XV. —Bees and Flowers Mutually Complementary - 247

XVI. —Bees as Fertilisers, Florists, and Fruit-producers 279

BEES & BEE-KEEPING.

INTRODUCTION.

The Hive Bee, from the wonders of its social

economy calling into play instincts as remarkable

as they are inexplicable, has for many ages com¬
manded the interest and the admiration of men;
and since it is at least the storer of saccharine sub¬
stances in a most convenient and delicious form,
which, before the general introduction of the sugar¬
cane, supplied in a unique manner a human want, it
has been, during many centuries, associated with
mankind by a bond of necessity second only,

perhaps, to that which linked our forefathers to the
cow, the sheep, and the horse. Although we are

less dependent upon its untiring industry, its sweet
product and tuneful hum are as grateful as ever;
while scientific investigation has shown to us that
the little labourer is a prodigy of wonders, of which
those of a hundred years since had not a suspicion ;
and, as revelation after revelation opens up before

BEES AND BEE-KEEPING.

us, our interest intensifies and our minds delight
themselves, as they ever must when properly
engaged in studying the works of creation. Very
much, however, that has passed current as accurate
and established, has not borne the test of recent
scrutiny ; and it will be in part the object of the
following pages to expose mistake, and supply its
place, where practicable, by truth. The departure of
fable will, however, never leave a void, since human
imaginings are always unequal to Nature's resources ;
so that here, as everywhere, “ fact is stranger than
fiction.” In treating far more completely than has
previously been attempted the anatomy and physio¬
logy of the insect which has made for itself by far
the largest place in literature—the sluggard-rebuking
ant not even excepted—the writer will be found
frequently to differ from the conclusions of others;
but never has he ventured so to do without the
most careful and scrupulous investigation, aided by
the most refined microscopical appliances. Again
and again he will, in obedience to truth, be forced
to show that many time-honoured statements have
originated, not in painstaking study, but in crude
and daring guessing, or in a carelessness of obser¬
vation almost equally blameworthy. On the other
hand, however, the pleasure will often fall to him of
pointing to the discoveries of such men as Siebold,
Leydig, and Schiemenz, amongst naturalists, as well as
to the achievements of the older apicultural worthies,
and the beneficial results of the energy and per¬
severance of many still amongst us, whose names
are familiar as household words.

INTRODUCTION.

A very large part of our matter will be in all
respects absolutely new, being the issue of researches,
dissections, and experiments, which have, in connection
with the practical work of the apiary, occupied
delightfully no inconsiderable fraction of many years
of a busy life. The anatomical, physiological, and
botanical illustrations, which, to a work like the
present, are as important as the text, have been in
every case, save one or two, drawn by the writer on
the wood, direct from his microscopical preparations,
or the objects in situ , as the case may be; and it is
hoped that they may form in themselves a contri¬
bution to the general knowledge of the subject,
which may advance apiculture a stage in the
direction of a true science. The accuracy of the
drawing may, it is believed, be relied upon; but,
notwithstanding earnest effort on the part of
draughtsman and engraver, producing results which
it is felt will not suffer by comparison, it must still be
confessed that the subtleties of Nature are in advance
of the refinements of Art, and that it has not been
possible in every case to secure full details. The
drawings are all to scale, which has usually been
given with the description. This fact will make the
illustrations of appliances peculiarly serviceable, since
all measurements can be readily ascertained, even
where they have not been stated.

Our title is a compound one, and our treatment
shall be complex. Practical bee-keeping is the out¬
come, and not the parent, of a scientific knowledge
of bees and their relations to the world about them.
Practical men have not made scientific apiculture;

BEES AND BEE-KEEPING.

but scientific men have given to practical ones not
only true methods, but reasons for their truth, and
so we logically place our scientific matter first, and
then look at our systems of operation in the light
thus gained. “ Practice makes perfect,” is but a
half truth. Practice, without intelligent insight, only
stereotypes; but practice, hand-in-hand with accurate
knowledge and observation, works out perfection. It
is our hope, then, not only to delight the student of
Nature by introducing to him beauties of structure,
wonders of adaptation, and minute refinements, to
which our conception is almost unequal, but to aid
to the full the bee-keeper, who can be charmed
through the pocket as well as the imagination.
Apiculture may be, and often is, profitable, ah,
very profitable, in the hands of those who would
not claim any scientific acquaintance with it; but
such from the teachings of others always adopt
scientific methods. While the course of events is
not unusual, deputed knowledge may be enough; but
management of the highest type — i.e., the most
remunerative kind, can only be arrived at by some¬
thing better than rule of thumb. In critical matters,
the best informed are the most trustworthy guides,
and knowledge, which appears to have little to do
with the practical side of the question, not infre¬
quently turns out to be the solver of an otherwise
unsolvable mystery, and the source of the best,
because truly scientific, method of procedure.

The ever-increasing zest attaching to apiculture,
the multiplication of bee-keepers, the competition of
dealers, and the ingenuity of inventors, has augmented

INTRODUCTION.

appliances in a surprising and somewhat bewildering
fashion. The endeavour to increase profits, and
make a market for honey, by saving the labour of
the bees and their owners, and by tempting the
purchasers of the results of the efforts of both, has
put before us a miscellany of articles, which our
ancestors would have regarded as fearfully and
wonderfully made, but which they never could have
supposed to have any relation to bee-keeping. It
will be our desire to do full justice to these, both by
explanation and illustration; but, at the same time,
by carefully elucidating principles, to so guide the
bee-keeper, that he may be well able to so select
both hives and appliances, that all his requirements
will be met by the smallest possible collection of
bee paraphernalia.

The increased attention which apiculture has
received during recent years, not only in our own
country, but on the Continents of Europe and America,
is due to a variety of causes. With us, some of these
are personal; and first in the roll of honour amongst
those who have with philanthropic ends attracted
attention to apiculture, must stand the name of the
Rev. Herbert R. Peel, whose very recent death all
must deplore, especially such as knew him so
intimately as the author. But the causes mainly are
of a more permanent kind than can be those that are
associated with the proverbial uncertainty of human
life; e.g., honey is a wholesome delicacy, which sugar
has supplanted, but not replaced ; so that this product
of the apiary is winning for itself anew a position
in our diet tables, and we are also beginning to re-

BEES AND BEE-KEEPING.

discover virtues in honey which had been well-
nigh forgotten. Old systems were clumsy and
uncertain, and yielded, at the best, but poor results.
Our modern plans give us complete mastery of our
bees, and enable us to obtain from five to ten times
the weight of honey from a single stock that the old
hands ever secured. Their honey in the comb was
generally stained, always irregular, and never to be
touched without leaving a sticky trail; ours, if we
know the art, is faultless in colour, flat as marble slabs,
and would not sully the daintiest glove. Theirs, when
“ drained,” or “ squeezed,” was often dirty and con¬
taminated by brood juices; ours, thrown out by the
extractor, is bright and clear, and of perfect purity.
It is no wonder, then, that purchasers increase and
apiculture is stimulated. There is a charm, too, in
modern bee-keeping, which never existed when the
hive was a sealed book and the bee supposed to
possess two points of interest only, and those at its
extremities —• its tongue and its sting — which had
nothing particular between them—to use the words
of a humorous writer—save “ skin and squash.”

The amateur, the naturalist, and the trader, alike
find more to delight and attract than was formerly
possible, while the general public are beginning to be
more alive to the advantages which honey possesses
as a food. Apiculture, then, has a raison d’etre
which assures its permanence, a pleasing thought to
those who know how much bees have to do with
securing for us a fruit crop, and fertilising many of
the plants cultivated by the farmer.

CHAPTER I.

WILD AND HIVE BEES.

The Position of the Hive Bee in the Animal World —
System of Classification—Family Apidse—Megachile
centuncularis—Humble Bees.

Nature, with a prodigality which bespeaks infinite
resources, has spread before the bewildered naturalist
between a quarter and half a million species of creatures
inhabiting land and sea. To marshal into system this
vast host, certain marked characteristics have at first
been laid hold of, so that all might be collected into
a few divisions. The classification of Huxley, which
I shall adopt, thus brings the whole animal world
under seven heads, denominated sub-kingdoms, because
life, in the widest sense, has been arranged under
kingdoms — the animal and vegetable. The second
sub-kingdom, Annulosa, embraces all those whose
bodies are definitely arranged in rings, including such
unlike creatures as house flies and leeches, so that
this sub-kingdom is naturally split into two parts—
Arthropoda and Anarthropodaf meaning those that

* apOpov, a joint, and Trou?,

foot, with the Greek privative av.

BEES AND BEE-KEEPING.

have and those that have not jointed feet. The fly and
leech thus part company, while, of course, our bee
takes its place in the first division ; but even here we
have wide diversities between creatures that cannot
claim kinship, such as butterflies, spiders, and lobsters,
the whole of which conform to the distinctions up to this.
point established. The A rthropoda are, therefore, sepa¬
rated into four classes, the first of which is Insecta —
insects having all certain well-marked peculiarities that
will appear in the sequel; but it is sufficiently exact
for our present purpose to now briefly state that the
Hive Bee is an insect because its frame is divided
by deep constrictions into three parts. First, the
head; second, the thorax or chest, to which are arti¬
culated or jointed the legs and wings; and third, the
abdomen.

Much as we have now narrowed our limits, this
definition still embraces a vast multitude of creatures
—comprehending moths, beetles, and flies—which
would appear to have little affinity with bees, and
so, for purposes of classification, other distinctions
are introduced, insects being separated into thirteen
orders, of which the Hymenoptera , or those carrying
four gauzy wings, includes not only bees, but also
wasps, ants, and some others. The Hymenoptera
being again parcelled out into families, distinct places
are found for the latter insects, while our favourites
appear amongst the Apidae, or long-tongued bees,
which, in company with the Andrenidae, or short-tongued
bees, comprise about 2000 distinct species, of which
212 are acknowledged natives of Britain, and these,
although differing greatly amongst themselves in size,

WILD AND HIVE BEES.

colouring, and habits, possess strong resemblances
in structure, suiting them all for honey and pollen
gatherers.

These pre-eminently useful little labourers forming
the families of the Apidse and A ndrenidse, as we
have already said, and of the merits of whose work
we cannot speak until we come to discuss their
relations to flowering plants, are far too much
strangers to bee-keepers. The big Humble is every¬
where recognised, and frequently its nest is not
unknown ; but the smaller solitary bees are not
certainly acquaintances of the ordinary bee-keeper,
notwithstanding his deep interest in their near
relative. It will be well here, therefore, to introduce
one or two for future identification, and these will
also serve for the purposes of illustration and com¬
parison.

Standing by a rose bush, we note the descent of
an insect, somewhat less than a honey bee, black-
backed, with reddish hairs on the thorax, and light
down upon its head and three first abdominal
segments. It poises itself a moment above a selected
leaf, and, settling, immediately commences cutting
with its mandibles, which act like a pair of scissors.
Quickly, a most regularly formed piece is detached,
which does not fall, heavy though it appears in
comparison with the size of the insect, for legs and
jaws continue to hold it, and away she flies towards
her nest.

I examined one of the latter recently, which had
been dug in the side of a quiet lane. The Megachile
centuncularis , for such is the name of this little

BEES AND BEE-KEEPING.

bee, had excavated a hole, at first perpendicularly,
and then horizontally, about 5m. in length. The
work of lining the tube with leaf had commenced
by cutting from some rose bush a circular piece,
curling this, and carrying it to the bottom of the
tube, and spreading it, without a wrinkle, into a saucer
form, to cover the end. Now the jaw scissors had
been set to snip out from the leaf-side spindle-
shaped pieces, which, brought one by one to the nest,
are applied to the wall at the bottom, and made
to overlap so cleverly that the earth is entirely
covered, while the serrations of one piece, worked
alternately in front of and behind the cut edge of
the next, hold all in exact position without any
cementing. We have now the representative of one
cell of ordinary honeycomb, and the analogy con¬
tinues in that the Megachile proceeds to her feeding-
ground amongst the thistles, from which she collects
pollen by hair brushes on her hind legs, whence
it is conveyed for temporary storing to feather-like
appendages on the under side of the abdomen;
honey is gathered by her tongue ; and thus fur¬
nished, she proceeds homewards to practise the art
of pudding making, for the two materials are kneaded
together, and increased in volume by repeated visits
to the thistles, until a stock of food, in all respects
resembling that used in the bee-hive, and sufficient
for one of her progeny, fills her leaf-lining nearly
to the top. Her first egg is now deposited, from
which, in due course, will issue the humble grub,
which, through Nature’s far-sightedness, with all its
humility, is still born to a competency. The cell

WILD AND HIVE BEES.

needs closing, while its cover is made to form the
floor of the next. Once more, then, the rose bush
must contribute perfect circles, for the cutting of
which no compasses are required. To the number
of four or five these are laid, one upon the other,
and pressed smoothly into position; the wall-lining
is added, a second pudding and egg provided, and
the processes repeated until five or six chambers
are complete, and the work of the little labourer
brought to a close. And now, strangely, the last
deposited egg is the first in the order of time to
hatch. The grub emerging does as a grub so
placed must: it consumes its pudding, and begins
to occupy the space its food previously filled. The
mother had accurately judged, if she could judge
at all, the needs of her son, for this grub is a
male. The pudding is gone, and he is satisfied, and
now begins to spin a cocoon, and then passes into
the chrysalis condition, and presently we have the
perfect male Megachile biting and pushing up the
leafy cover, and escaping into the sunlight of a new
life. By his emergence, he has opened up the way
for his brother, and he in turn will remove all im¬
pediment to the escape of the sisters below. Thus
the community of young Megachile is provided. The
old ones are gone, but the race lives. Their marriage
bells are rung while the autumn sun is shining; the
males die, the females seek screening from the chilly
blasts which must blow before their work of nest
building can commence, and so the circle is com¬
pleted.

How unlike, and yet how very like, all this to the

BEES AND BEE-KEEPING.

Hive Bee. As we become acquainted with the latter,
we shall see that the mother Megachile is queen
and worker combined ; the male, the short-lived
drone. The cell, its sealing, the food, the egg, the
tongue, the hair brushes, the abdomen of the two
insects, counterparts of one another in each case. All
Nature is one, and the student of the Hive Bee is
unwise, and self-deprived of the knowledge of much
that is marvellous and delightful, if he altogether
neglect all members of the family Apidse save its head
and most perfect representative.

Fig. 1.—The Humble Bee (Bombus).

The Humble Bees, or Bombi (Fig. i), come nearest
to our Hive Bees in that they are semi-social, living
in companies during the summer, the queen pass¬
ing the winter in solitude. The big downy and noisy
insect that visits our gardens in the spring is a
mother Bombus , that spent her honeymoon the pre¬
vious autumn, in like fashion with the Megachile ,
and subsequently sought out for herself some narrow
retreat in which to hybernate; but, so soon as the
sallows yield their pollen, she is abroad and preparing
for the progeny by which she is presently to be sur¬
rounded. Different species have different habits, but

WILD AND HIVE BEES. 13

in every case the hybernated mother commences a
nest alone ; e.g. y the Bombus muscorum, known by
its light and dark brown hairs, establishes itself not
infrequently in the middle of fields, taking care that
the spot selected is in the neighbourhood of abund¬
ance of flowering plants. A vaulted roof is formed
of cleverly entangled pieces of moss, plastered beneath
by a layer of greyish wax, and so rains, which would
pass the moss, are effectually kept out. Pollen and
honey are collected in pellets, eggs laid, and so, in
due course, workers produced; for, as her children
gather about her, the mother leaves to them the
duties of nest extension, cell construction, and food
collection, and, confining herself to ovipositing, becomes
a stay-at-home, and a very close representative of
the so-called queen of the bee-hive.

Later in the season, instead of workers, which are
much smaller than the queen, a Bombus, a size between
the two, begins to make its appearance in the nest;
this is the male, and now, soon, creatures as large as
the original mother are added to the colony. These,
the true females, mate as we have hinted, and alone
survive the rigour of winter to be the instruments for
continuing the race. To those conversant with Hive
Bees, the closeness of the analogies between the two
insects last mentioned, will suggest themselves; but
they will become evident to all as we study the next
chapter. Amidst the analogies, however, there are
differences, and so the family of British Apidse are
marked off into nineteen genera, the typical genus
being Apis, in which the Hive Bee finds its place.
In this genus, there is but one British species, mellifica,

i 4

BEES AND BEE-KEEPING.

although some others are cultivated as imported bees,
and notably Apis ligustica, or the Ligurian bee; so
that the whole classification of the subject of our
study would take the form now given :

Kingdom .
Sub-Kingdom
Division.
Class. . .

Order
Family
Genus
Species .

Animal.

Annulosa.

Arthropoda.

Insecta.

Hymenoptera.

Apidae.

Apis.

Mellifica.

CHAPTER II.

ECONOMY OF HIVE BEE.

Gathering Bees — Comb—Pollen and Honey — Eggs ,
Larvae , and Pupae — Foragers and Nurses —

Skin and Bowel Moulting and Cocoon Spinning
— Drones and Queens — Swarming and Comb
Building.

In order that we may possess an intelligent under¬
standing of the need and suitability of the various
complex parts and organs proper to Hive Bees, and
which we are about to consider in detail, it will be
necessary for us to pass in review a general outline
of the economy of the hive, noting at present only
the salient points. We will imagine that our study
is undertaken on a fine summer day, and that the
hive we have at command is in a normal, healthy,
and prosperous condition, and such an one also as to
afford us every facility for examination. As we stand
before its entrance, bees in quick succession make
their appearance at the hive door, and in such
haste as seems to indicate that they are impressed
with the importance of their mission, for they are no
sooner well visible than they are away in some
definite direction; but others are returning, and these,

BEES AND BEE-KEEPING.

settling, in a great number of instances, show us
that they are carrying on their hind legs relatively
large masses of coloured material, which is most
generally some shade of yellow or orange, although
crimson, green, and even black, may be seen. This
material, considered by the ancients to be wax,
and called by Reaumur himself la cire brute
(crude wax), we shall, in due course, learn to be
pollen, which has been gathered from the anthers, or
male organs of blooms, by a most beautiful set of
apparatus, to be hereafter examined. Opening our hive

by the removal of the top cover, so as to expose our
stock (as we commonly denominate a colony of bees
in an established condition), and in order that we may
learn the behaviour of those that are returning from
their aerial voyages, we find it filled with combs,
each one of which is a tolerably flat slab, about iin.
in thickness, fixed in, and mainly hanging from, the
upper side or top bar of such a frame as is shown at
Fig. 2. These frames are so placed and arranged
that each may be easily lifted out with its attached
comb, which has, in turn, its face £in. distant from

ECONOMY OF HIVE BEE.

its fellow on either side. These interspaces are well
filled by bees, but very few of which disturb them¬
selves on our account; nor need we be disturbed
on theirs if, with precaution, we lift out one of these
filled frames for inspection, the bees retaining their

Fig. 3.—Honeycomb (Natural Size).

Queen Cell, from which Queen has hatched, showing Lid; B, Queen Cell
torn open ; C, Queen Cell cut down ; D, Drone Grub ; E, Drone Cell,
partly sealed ; F, Drone Cells, sealed ; G, Worker Cells, sealed ; H,
Old Queen Cell; I, Sealed Honey; K, Fresh Pollen Masses ; L, Cells
nearly filled by Pollen ; M, Aborted Queen Cell on Face of Comb ; N
Bee biting its way out of Cell; O, Eggs and Larva* in various conditions.

sition, and in large part continuing their work
though nothing particular had happened.

A.s we now cast our eyes over the comb, delicately
d perfectly modelled in wax, we discover that

BEES AND BEE-KEEPING.

it is made up of a number of chambers (technically
cells), nearly all of which are exactly hexagonal

in cross section, and most of which are precisely
one-fifth of an inch between the parallel sides.
Some of these (L, Fig. 3) are nearly filled by an
opaque, dough-like looking body, which we recog¬

nise at once, both by colour and consistency, as
being that very pollen, packed away, which we saw
being carried into the hive on the hind legs of

the returning bees. One of the latter, still loaded,

marches before us, occasionally sharply agitating her
body (for these untiring workers are ladies) ; and,
as we look, she curls herself over a cell, and, by
a process singularly beautiful, which we are not
yet in a position to understand, she thrusts off one
of these lumps into it, and then, by a second twist,
the other, either immediately leaving them, as at K,
Fig. 3, or else butting them down with her head into
a pancake for future use. But her cargo is as yet
only half discharged; and now, seeking another cell,
either empty or containing some honey, she inserts her
tongue, and returns from an interior cavity of her body,
called the honey sac (h s, Plate I.), the sweet fluid she
has collected from the nectaries of flowers. As duty
and pleasure are synonyms with a bee, she at once hies
away, in order that, ere long, she may yet again add
to the riches of the community for which she lives to
labour. And, we ask, why this anxiety to carry
home both pollen and honey ?—some of the latter
standing before us in considerable quantity, beautifully
covered by air-tight, either white or yellowish, caps
of wax, seen in Fig. 3, at I. A reply is soon

ECONOMY OF HIVE BEE.

furnished, as we notice, at the bottoms of numbers
of the cells, whitish tiny legless grubs (O, Fig. 3),
evidently incapable of seeking their own food in any
way. This is brought to them by the younger bees
of the stock, which do not normally fly abroad, but
which make the helpless larvae (grubs) the especial
objects of their care, elaborating for them the two
kinds of “ pap,” which form their sole nourishment,
by a process respecting which great errors have
been propagated. We shall have much to say
about it presently. The materials required for the
somewhat circuitous elaboration of the given food are
honey, pollen, and water, which last, if need be, is
brought home in quantity, the former two being
placed in the store cells, as we now understand,
by the foragers, the name commonly given to the
flying bees, while the feeding bees are very ap¬
propriately called nurses, although there is no actual
distinction between them, as some former writers
thought. Growing older, the nurses turn to foraging,
but they do this in consequence of a glandular
change coming on with age, which makes nurse
work unsuitable; but more of this hereafter. This
pap may be seen, in appearance like arrowroot made
with water, surrounding the bodies of the grubs (see
FL, Fig. 4). They partially float in it, and, besides
absorbing it by the mouth, are commonly supposed to
take it in by that part of the skin which is submerged ;
but it is not correct, as stated by Cook, for reasons
presently given, to say that the food is “ all capable
of nourishment, and thus all assimilated.”

These model nurses are ever perambulating the

BEES AND BEE-KEEPING.

combs, and, in the darkness of the hive, so examine
the contents of every cell, by exploring it with
their thread-like antennae—which are most sensitive
organs, placed between the eyes, and well seen in
many subsequent illustrations, especially Plate II.—that
no grub escapes due attention, and food follows
close upon appetite, although, in a strong colony, often
as many as 12,000 larvae will need pretty frequent
visitation. The larva, or grub, grows apace, but not
without experiencing a difficulty to which the
human family is, in some sort, subject in the period
of youth. Its coat is inelastic, and does not grow
with the wearer, so that it soon, fitting badly, has to
be thrown off ; but, happily, in the case of the larva,
a new and larger one has already been formed be¬
neath it, and the discarded garment, more delicate
than gossamer, is pushed to the bottom of the cell.
It would be singular, were it not for the abounding
errors of bee literature, that Reaumur and Huber
have asserted (followed by many others with a uni¬
formity which is not the outcome of investigation)
that the bee larva does not change its skin, but only
grows larger. A little patient looking would have
found the old and ruptured pellicles, and so pretty
conclusively have settled the question. In like
manner to the first, moult succeeds moult, to the
probable number of six, when, after about four days’
feeding, the well-nourished creature, loaded with
fat, lies at the lower part of the cell curled up,
as one is seen to do near H, Fig. 3. At this
time, its weight is scarcely less than double that of
the "bee into which its natural transformations will

ECONOMY OF HIVE BEE.

by-and-by convert it. No more food is supplied,
and the period for cocoon spinning approaches. The
silken threads forming this {co, Fig. 4) are produced
by a fluid yielded by a gland (Fig. 15), which
re-appears in the adult bee. This fluid escapes by
an aperture in the lip, and very quickly hardens
into what may be described as bee silk. Before the
cocoon can be built, a cover, technically called
sealing, is put over the larva by its nurses, that
now bid it farewell. These covers are seen in num¬
bers at G, Fig. 3 ; they are pervious to the air, are
made of pollen and wax, and are more convex and
regular in form than those sealing in the honey
(I, Fig. 3) ; and, behind them, a series of most
wonderful and bewildering changes occur; but, ere
they can commence, a preliminary step is necessary,
which seems to have altogether escaped the attention
of both scientific and practical writers. The food
given to the larva, especially during the latter part
of the growing period, contains much pollen, the cases
of the grains of which consist of a substance called
cellulose, which is perfectly incapable of digestion.
These cases, with other refuse matters, collect in
quantity within the bowel, which becomes distended,
since it has no opening {mb, Fig. .13). The im¬
prisoned larva, having little more than enough room for
turning, must be freed of these objectionable residua;
but Nature is equal to the difficulty, accomplishing
all in a manner commanding our admiration—and here
we can but outline, reserving a fuller explanation till
we consider the structure of comb. In a word, the
larva turns its head upon its stomach, and pushes

BEES AND BEE-KEEPING.

the former towards the base of the cell until its
position is reversed, the tail being outwards, and,
thus placed, it laps up all residue of food, especially
from its old clothes previously referred to, until they
are dried, and practically occupy no space. It now
throws up its stomach and bowel, with all their con¬
tents, and without detaching them from its outer
skin, which is moulted as before, but, in this instance,
to be pressed against the cell, so as to form for it
an interior lining. The dejectamenta of the bowel in

Fig. 4.—Larva and Chrysalis (Magnified Four Times).

SL, Spinning Larva ; N, Nymph or Chrysalis ; FL, Feeding Larva; co, Cocoon;
sp, Spiracles ; t, Tongue; m, Mandible ; an, Antenna; to, Wing; ce, Compound
Eye ; e, Excrement; ex, Exuvium.

this way lie between the cast skin and cell wall (as
seen at e, Fig. 4), and so the larva remains abso¬
lutely unsoiled. It now turns its head and resumes
its old position, joining its cocoon to the edges of its
last cast skin, so that its habitation is relined, it is
cleansed, and air can still pass to it through the
imperceptible openings left by the bees in the seal¬
ing. This point is of radical importance, since
breathing is carried on pretty rapidly during the

ECONOMY OF HIVE BEE.

latter part of its subsequent transformations, the
absorbed oxygen permitting then of a production of
heat, and causing also considerable diminution in
weight. Having thus put in order the cell con¬
taining it, the larva remains for some little time in a
condition of quiescence, and now, under the new
name of chrysalis, pupa, or nymph (N, Fig. 4), enters
upon the sequence of transformations, all slowly and
quietly effected, which end in converting it into a
new creature. Constrictions occur, and rings or
segments vanish, until the body becomes head, thorax,
and abdomen. As it lies upon its back, prominences
begin to show themselves, which become more and
more pronounced, until, at last, they sufficiently assume
the form of legs to be recognised; these are six in
number and are much more than organs of loco¬
motion, as they bear, curiously disposed upon their
many joints, a whole set of tools singularly varied in
modelling and application ; a tongue, too ( t , Fig. 4),
replete with wonders, and lying stretched along above
the body, begins to be seen ; and then, drawn round
from the back of the thorax, like a girl's cloak which
she has allowed to slip from her shoulders, are gauzy
but many-folded extensions, which hereafter become the
beautiful instruments of flight {w, Fig. 4). But these
external changes, marked though they be, are trans¬
cended by the wonder of the progressing interior modi¬
fications and developments. The nerve system is recast
and enlarged, the digestive apparatus changed, an
entirely new set of muscles and tendons brought into
existence; glandular structures make their appearance,
breathing tubes or trachea in untold number come into

BEES AND BEE-KEEPING.

being, and, in short, an organisation is built up which
has baffled, and is still in great part baffling, our
highest powers of research. That which was the
blind grub, living in darkness, is soon to be the
active bee, rejoicing in the sunbeam, attracted both
by the perfume and the colour of flowers, and so
organs of sense are being prepared for it, the struc¬
ture of which we cannot yet stay to consider;
the antennae are developing, and, at the sides
of the head, dark brown spots are indicating the
position of the future compound eyes. In some¬
thing more than twelve days from the time of
sealing, the transformations are complete, and a
pellicle, delicate as cobweb, is rolled from every part
of the frame, and pushed downwards to the base of
the cell {ex, Fig. 4), where we soon may be at
liberty to find it, for now a creature, lacking in
nothing that its subsequent duties will require, bites
at the door of its prison-house, into which it soon
carves a long, curved slit, as seen in three or four
cases (Fig. 3) ; and then, by a push, it makes way
for its emergence, the head is advanced as at N, and
a pale but perfect bee walks into view. Its down,
like that of the recently-hatched chick, adheres, but
soon it will dry and preen itself, and in twenty-four
hours we shall have our nurse already entering upon her
duties to spend and be spent, in order that she render
to others those very attentions she has herself received.

But we ask, Whence the grubs whose history we
have so far examined ? and now, in searching, we
discover, on one of the combs, an insect—commonly
but very erroneously called the queen (Fig. 5, b ), for

ECONOMY OF HIVE BEE.

she in no sense governs—longer in body than the
worker (Fig. 5, a), and really differently formed in
every part, and possessing most active and curious
egg organs, called ovaries, which are capable of
yielding a prodigious quantity of eggs. As we watch
this queen slowly progressing, with a number of
workers about her, touching her continually with
their antennae, and backing out of the way so as not
to impede her movements, she dips her head very
deeply into a cell, and, having satisfied herself that
it is empty, she advances a step, holds on to the
edges of the comb, principally by her second and

third pairs of legs, and, curling her abdomen, inserts
it into the examined cell, until it is almost entirely
hidden. A moment of apparent stillness; she recom¬
mences her walk, her abdomen straightens as it rises
from its hiding place, and we immediately see that
she has left behind a tiny long and narrow pearly-
white egg, fixed by one end to the cell bottom. The
queen quickly repeats the operation, the neigh¬
bouring nurses being always ready to offer food.
Their attentions are, as we can easily see, needful,
but many writers have given the echo to a medieval
fancy by stating that she is ever surrounded by a

BEES AND BEE-KEEPING.

circle of dutiful subjects, reverently watching her
movements, and liable to instant banishment upon
any neglect of duty; these it was once the fashion
to compare to the twelve Apostles, and, to make
the ridiculous suggestion complete, their number
was said to be invariably twelve. If all this were
true, beginners in bee-keeping would not find the
difficulty in discovering a queen which they some¬
times experience. But to resume. The egg contains a
germ, which, kept warm by the native heat of the
colony, and fed by abundance of yolk, will develop
into a grub, which, in some instances, frees itself from
the egg case by struggling into the first quantities of
food put into its cell by the nurse bees, the very
condition in which we just now made its acquaint¬
ance.

We have already learnt that the worker bees are
female, but they are sexually aborted, and normally in¬
capable of laying eggs. The queen, or mother, on the
contrary, is fully developed, and her capacity for egg-
production is immense, a good queen being able to
furnish to the cells an average of two eggs per minute
for weeks in succession. A new question now arises,
Whence the queen ? and we are brought face to face
with a difficulty which even yet we may not have fully
surmounted, although, in a later chapter, I hope to
give a relatively more satisfactory answer than has
yet been attempted. The queen, in short, is pro¬
duced from an egg in all respects identical with the
eggs which furnish the workers. The difference is
brought about by a change of treatment to the grub
on the part of the nurses. When a queen is to be

ECONOMY OF HIVE BEE.

2 7

produced, a cell of large size and extraordinary form
is constructed (A and B, Fig. 3), and, by special
feeding of its occupant, instead of a worker, a queen
is evolved. She, being a female, needs a mate, and
such is found in the drone (c, Fig. 5), or male, which
has a very complicated structure, that must be duly
considered under its proper head. The drones are pro¬
duced in larger cells than the workers, so that their
more rotund forms may be accommodated. Their cells
are a quarter of an inch in diameter, and are seen
over D, E, F, Fig. 3, and may always be recognised
when they contain sealed brood (the name for inclosed
larvae) by the very convex forms of their cappings. The
eggs to provide these males are also laid by the queen,
and are, whilst in her ovaries, absolutely like those
that furnish both queens and workers. When, however,
the latter are to be evolved, by a somewhat compli¬
cated act occurring in the body of the queen just
before the egg is deposited, fertilisation takes place
by the addition of material originally received from
the drone. When drones are to be produced, this
addition is withheld and the eggs are laid unim¬
pregnated— i.e., drones have a mother, but no father,
a question, the examination of which, with the
anatomy of the parts involved, will be fully explained
and illustrated as we proceed with our task..

The tremendous fecundity of the queen, in favour¬
able conditions, so multiplies the number of bees, that
a division of the community becomes necessary,
beside which, these wondrous little animals have an
essential and deeply-rooted colonising instinct, upon
obedience to which they often insist with singular

BEES AND BEE-KEEPING.

pertinacity. Since a queen is essential to the ex¬
istence of a stock, as she alone can produce eggs,
a new queen, under these circumstances, must be
produced, and so bees form queen cells, and previously
bring forward drones. The old mother departs with
the superabundance of the population. A queen,
matured soon after her migration, occupies her place
after having consorted with a drone, so as to secure
the honours of maternity.

Such, in few words, is swarming. The swarm
needs powers we have not yet considered. Its new
house requires furnishing, and, to compass this, first
wax is secreted from the bodies of the workers, and
then, by an architecture which is rarely, if at all,
exceeded in beauty and adaptation even in the
insect world, combs are built of dainty purity and
almost mathematical exactitude (“almost” is here
said advisedly), and so a place is given, as the
cells multiply, for the eggs of the accompanying
mother and for the incoming riches brought home
by the never weary foragers ; and if weather be favour¬
able, or, what is even better at this particular point,
the bee-keeper intelligent and attentive, our swarm
quickly passes into a stock, and will yield us all the
interesting points which have as yet occupied our
attention. It is now clear that the mysteries of the
economy of the hive, the varied instincts brought
into exercise, and the wondrously complicated and
delicately beautiful organisations of the little labourers
making their purposeful lives a possibility, will give
much occupation during succeeding chapters.

CHAPTER III.

GENERAL STRUCTURE.

External Skeleton — Chitine—Hairs and their Uses —
Breathing Apparatus : Spiracles and Tracheae ;
Air Sacs—Circulating System—Dorsal Vessel —
Pericardial Cavity—Blood of Bee — Peritracheal
Circulation — Muscular Fibres , and Methods of
Movement—Muscles of the Java—The Minute in
Nature.

OUR bee is now before us, for we have witnessed
the laying of the egg, the growth of the larva, the
development of the chrysalis, and the initial life of
the imago, and, as we pursue our study of the
intricacies now awaiting us, we shall find it more
convenient to treat under separate heads the nerve
system, the digestive apparatus and glandular systems,
the external organs of sense and locomotion, with
those special parts that distinguish queens, workers,
and drones ; but, at the same time, it will do us good
service first so to examine the general structure that
we shall have a grasp, as a whole, of the wondrous
mechanism we desire to understand. Let us begin
with the external framework, premising that bees,
in common with all insects, have formed on every

BEES AND BEE-KEEPING.

part of their bodies, by a layer of secreting cells,
called the hypodermis, an external skeleton, com¬
posed of a remarkable substance, to which the name
Chitine has been given. Chitine is capable of
being moulded into almost every conceivable shape
and appearance. It forms the hard back of the
repulsive cockroach, the beautiful scale-like feathers
of the gaudy butterfly, the delicate membrane which
supports the lace-wing in mid air, the transparent
cornea covering the eyes of all insects, the almost
impalpable films cast by the moulting larvae, and the
black and yellow rings of our native and imported
bees, besides internal braces, tendons, membranes, and
ducts innumerable. The external skeleton, hard for the
most part, and varied in thickness in beautiful adap¬
tation to the strain to which it may be exposed, gives
persistency of form to the little wearer; but it needs,
wherever movement is necessary, to have delicate
extensions joining the edges of its unyielding plates.
This we may understand by examining the legs of
a lobster or crab, furnished, like those of the bee,
with a shelly case, but so large that no magnifying-
glass is required. Here we see that the thick coat
is reduced to a thin and easily creased membrane,
where, by flexion, one part is made to pass over the
other. Likewise, in the antennae of the bee [a, Plate II.),
the insertion into the head, by a sort of ball and
socket joint, covered by chitine so thin and transparent
that nerves may be seen through it, admits of the
varied movements proper to this instrument of inter¬
communication ; for it is hardly too much to say that, by
means of the antennae, the intelligent little creatures talk.

GENERAL STRUCTURE.

Again, almost every part of the body is covered
by hairs, the form, structure, direction, and position
of which, to the very smallest, have a meaning. These
are also formed of chitine, and framed for varied
uses. The external skeleton, mainly protective in
character, is not sensitive, and so a large propor¬
tion of these appendages are curiously formed (as at
C, D, E, and H, Fig. 24), with a bulb at the base,
to accommodate a nerve end, by the presence of
which they become, in each individual instance, truly
organs of touch. Beside this, they act as clothing,
the thoracic and abdominal pubescence, or fluff, aiding
in retaining heat, and give protection as the stiff,
straight hairs of the eyes (Plate II.), whilst some act
as brushes for cleaning ( eb , C, Plate V.) ; others are
thin and webbed, for holding pollen grains (as I,
Fig. 24) ; whilst, by varied modifications, others again
act as graspers, sieves, piercers, or mechanical stops
to limit excessive movement. Possibly, the hairs are
not exclusively utilitarian, since those on the dorsal
part of the abdominal rings would appear to be
intended mainly as a decoration.

Whilst carefully scrutinising a worker and a drone
by fhe aid of a hand magnifier, or watchmaker’s eye¬
glass—and every intelligent bee-keeper should at least
possess some such apparatus — we note that the
abdomen of the worker, like that of the queen, is
surrounded by six belts of chitine, each being made
of two plates—one, larger, on the back (the dorsal
plate), overlapping the second, smaller, or ventral
plate, which is applied to the lower side of the body.
This arrangement is well shown in the chrysalis

BEES AND BEE-KEEPING.

(N, Fig. 4), or in the cross section of imago (Fig. 8).
The drone is similarly formed, but has seven belts
or rings. These, in both cases, if the specimens are
living, are continually slipping in and out upon
each other like the joints of a telescope, their attach¬

es Air Sac, showing Plaiting; b, Spiral Threads of Trachea ; c, Interior, showing
threads; d. Single Thread from Small Trachea; e, Fine Tracheae.

ments being made by delicate membranes, which
admit of free movement (abdomen, Plate I.). As
these slide backwards and forwards, we catch sight
of depressions that are hidden from view when the
abdomen is fully drawn in, one occurring near each

GENERAL STRUCTURE.

end ,of each dorsal plate save the first. Microscopic
examination reveals that we have here openings,
denominated spiracles ( sp , Fig. 8), with strange com¬
plications, leading into internal tubes, called tracheae
(Fig. 6), forming the breathing apparatus, and which
divide and sub-divide, after the manner of a fibrous
root in the soil, until they are found in countless
number in every part.

All animals require oxygen. In those above the
Annulosa (page 7), the blood is carried either into
lungs or gills by means of vessels, when it appropri¬
ates oxygen, which, by the circulation, it distributes.
In insects, with a local exception noticed later in
the chapter, there is no system of blood vessels, so
oxygen, as a part of the air, is taken direct from
the before-mentioned spiracles, through the tracheae,
into all muscles, glands, and organs of the body,
not even excepting the wings. As the abdomen
is extended and contracted, as is constantly done by
the bee, air is drawn into, and then expelled from,
these apertures in the sides, precisely as in our own
breathing from the mouth. Should an unlucky fly,
through not sufficiently controlling his passions and
appetites, tumble into the milk, and be saved from a
tragical fate by being lifted on to the table-cover, he
immediately commences energetically grooming his
body with his legs, not because he is especially anxious
about his personal appearance, but because here the
milk is closing his spiracles, and actually choking him.

The tracheae consist of an external and internal
membrane, between which run spiral threads, highly
elastic in character, that prevent the closing of the

BEES AND BEE-KEEPING.

tube by any bending of the body of the insect, just
as the spiral wire within indiarubber gas-piping secures
a constant flow of gas, in spite of any twisting of
the pipe itself. The embryology of insects has shown
that the tracheae are developed by invagination (a turn¬
ing inwards) of the outside skin (precisely as the
bowel is formed in the larva, see Fig. 13), and that, at
the time of moulting, the tubes in the neighbourhood of
the spiracles are cast off. That this is true in the bee
is easily proved by those having a microscope of even

moderate capability. Lifting from a cell a half-grown
larva, a little transparent mass will be observed upon
the centre of the cell base, which mass to some extent
filled the cavity formed beneath the body as the grub
lay head and tail together. This is found to contain
one or more cast skins, which carry with them the
covers of the spiracles. The investing membrane
(Fig. 7) of the contiguous tubes is withdrawn, while
the tiny hairs and scales of the body also lose a layer
as we see by the illustration. It is difficult to under¬
stand how the extremely thin lining of the tracheae is

GENERAL STRUCTURE.

removed, but the fact is evident. From the invagina¬
tion aforesaid, it follows that the layer which is
outside in the skeleton is the inner, or lining one, in
the tracheae, while the hypodermis, which originates
the chitinous coat (as has already been stated), and
lies, of course, beneath it, has its representative out¬
side the breathing tubes. The spiral thread is pro¬
duced by the lining membrane, or internal cuticle,
forming a chitinous thickening, in a spiral line, which
is never continuous for more than four or five turns.
Just before one thread terminates a new one starts,
to be in like manner followed by another. The tubes
are only capable of slight extension, and, when
unduly stretched, the membrane ruptures, and the
spiral is drawn out singly (as at d, Fig. 6), or a
band of four or five threads will separate for a few
turns (as at b ). The slenderness of the smallest of
these tubes, which have neither interior cuticle nor
spiral thread, is as remarkable as their number, and
the microscope, even at its best, is barely able to
trace out their terminations. Of such, a bundle
containing a quarter of a million, would scarcely
exceed in bulk an ordinary human hair.

In bees, as in all actively flying insects, the
tracheae are accompanied by large air sacs (a,
Fig. 6), which are developed in the same manner as
the tubes themselves, but carrying scattered venations
instead of spiral thickenings of the membrane. In
the larval state of comparative inactivity no aerial
sacs exist, but they are brought into being during
the chrysalis changes. These air sacs have much to
do with flight, in a way to be explained when we treat

BEES AND BEE-KEEPING.

of the wings, while their forms ate such, that no draw¬
ing professing to embrace them all could do more
than give an inadequate idea. The main ones, in
the worker and drone, lie in the anterior part of
the abdomen, on each side, communicating with the
spiracles ; but in the queen they are greatly reduced,
to give room for the ovaries.

The spiracles are simple in the larva (. sp , Fig. 4),
and twenty-two in number (on each side ten well-
developed and one rudimentary, the latter vanishing
altogether before the last moult. The oft-repeated
statement that they are eighteen in all is an error).
In the adult, they are more complex, capable of
voluntary closing, and so arranged that foreign
bodies cannot accidentally - enter, while their number
is only fourteen—five on each side of the abdomen,
and one behind the insertion of each wing.
During the period of pupa-hood some of the rings
possessed by the larva disappear, while the spiracles
they carried vanish. Hence, the adult bee has
fewer than the grub,, whence it came. In the
drone, the spiracles are much stronger and larger,
and so more easily studied than in the worker.. They
are furnished with an apparatus to add to the noise
of the insect’s flight, which will be more fully noticed
by-and-by, are surrounded by delicate protecting
hairs, to save them from dust, and number sixteen,
in consequence of the drone having an additional
abdominal segment. The normal respirations of the
bee, when at rest, varying from twenty to fifty per
minute, are much influenced by external temperature,
by the activity of the stock, and by the amount of

GENERAL STRUCTURE.

heat it may be necessary to maintain so as to best
suit the condition of the . brood chamber.

Although insects, and bees in the number, have no
general system of blood vessels, as I just now said,
they still have a beautiful apparatus by which their
fluids are continiially carried round and made in
purity to visit for nourishment and renewal every
part of the body. Their heart, or blood pump, is
called, on account of its position, the dorsal vessel,
for it runs as a complex tube {dv, Plate L, and Fig.
8) along the back, almost immediately beneath the
external skeleton. This heart may be seen in action
in almost every caterpillar, where the opening and
closing of the ventricles, as they are called (z>, Plate I.),
can be watched through the semi-transparent skin.
If we are fortunate enough to possess a microscope,
we may very easily see the pulsations far more
beautifully in the tiniest of the larvae. Remove
from its cell with a blunt needle the smallest to
be found, place it on a glass slip, add a drop of
water, and, with gentleness, a thin cover glass,
when the transparent larva will show', with an inch
objective, many wonders beside its spiracles and
tracheae, digestive tube, and nerve system, with the
dorsal vessel continuing for some time to gently
pulsate. Without a microscope, a little manual
dexterity will make the movements of the heart
visible in the adult bee. If one accidentally injured
is not at hand, a victim to science must be decapi¬
tated, and then opened on the under side of the
abdomen, so as to remove the stoipach and expose
the mere back shell seen from within, as Fig. 8 will

BEES AND BEE-KEEPING.

make clear, when sharp eyes, or weak ones with a
lens, will detect rhythmic throbbings, continuing long,
and moving from behind forwards, driving the blood
towards the head, much as water rises through the
throat of a drinking horse. The walls of this heart
consist of three layers—an internal cuticle, a central
layer of muscular fibres, zoo^in. in diameter, and an
outer coat of connective tissue. In the worker and
queen, the dorsal vessel has five ventricles, or con¬
tractile chambers, corresponding to the five spiracles
on each side. As it nears the thorax, the muscular
and internal layer now formed into a conducting
tube, bends upon itself three or four times from side
to side (Plate I.), by which I imagine the rhythmic
beats are converted into a steady and equal dis¬
charge of blood in the head beyond, where the tube
opens near to the brain. The vitalising fluid returns
by soakage through the body to the posterior part,
where it re-enters the dorsal vessel. The ventricles
are in valvular communication, while each one has
on its sides two openings ( dv , Fig. 8), so contrived
that, as the muscular coat is causing a ventricle to
dilate, blood enters by them, the valve in front at
this time closing, as Plate I. will explain. When
contraction begins, an internal fold of the wall of
the ventricle closes the side apertures, and drives
the ' blood through the communication into the
ventricle in front, and in this manner the forward
stream is maintained. The dorsal vessel is braced to
the dermal skeleton by surrounding muscles, while
beneath runs an extension of muscular plates
(d, Fig. 8) of most involved character, forming a

GENERAL STRUCTURE.

horizontal diaphragm or division wall separating the
abdomen into two very unequal parts, the larger of
which is below.

This diaphragm* in contracting increases the upper
cavity and diminishes the lower, and so pressing to¬
gether the viscera, drives from them blood, which
now enters the heart chamber or pericardial cavity
( pc , Fig. 8), by apertures in the diaphragm itself.

Fig. 8.—Cross Section of Abdomen of Worker Bee (Magnified Bight times).

dv, Dorsal Vessel; d, Diaphragm; pc, Pericardial Cavity; sp. Spiracle; tr,
Tracheae ; ts, Tracheal Sac ; at, Stomach ; n, Nerve ; ga , Ganglion.

The dorsal vessel presents many microscopical
curiosities ; it rests upon a cushion of pericardial
cells, with singular nuclei, and which sometimes
send extensions either into the outer layer of the
heart or the diaphragm. We also find here lobes
of fatty bodies (corps graisseux ), containing here and
there the cellules enclavees , or separate cells of
Graber, of yellow colour, with a single nucleus, and
which resists the action of acids and alkalies, and,
beside, multitudes of nerve filaments, and some exceed¬
ingly fine ramifications of the tracheal system.

* This diaphragm has been investigated by Graber (see “ Archiv fur
Anat. microscop de Schnltze,” vol. ix., p. 129).

BEES AND BEE-KEEPING.

The blood of the bee is colourless, and contains
but few corpuscles, which are always white, and
carry a nucleus surrounded by granular matter, and
have the wonderful though not unusual quality of
constantly changing their outline, whence they are
called amoeboid. At one moment they will be round,
but slowly they become ellipsoid, and then, perhaps,
an irregular boat shape, or even star-formed..

Our subject is so vast that space can hardly be
spared for the discussion of exploded theories; but
some mention must be made of the so-called “peri¬
tracheal circulation,” a pet notion with M. Emile
Blanchard. It was supposed that the blood was
carried along the tracheae, between their two walls.
The idea was based upon a misunderstood experi¬
ment, and the microscope gave no countenance to
it; it may now be regarded as beyond resuscitation.
The coup de grace was administered in Graber’s
explanation of the functions of the diaphragm, which
has removed a great difficulty, as it is now seen that
the blood in the pericardial cavity is enriched with
oxygen, by the numerous fine tracheae there placed,
and sent in best condition into the dorsal vessel to
supply first the brain, and then, in turn, every part.

The muscular system, by which all movements are
brought about, depends for its action upon nerve,
which induces a contraction, bringing nearer together
the parts attached to the extremities of the fibres
building up the muscle. The individual fibres, parts
of two of which are represented in Fig. 9, are very
varied in size in bees. The largest with which I am
acquainted are those forming the powerful muscles

GENERAL STRUCTURE.

41-

enabling the drone to contract the abdomen so as
to produce the expulsive act. One of these fibres,
where shortened and thickened (as at a, Fig. 9),
may measure ^th of an inch in diameter, while
the relaxed portion of the fibre is about rj^th. If
this muscle be skilfully and quickly removed, and
placed either in the fluids of the animal or in a
little weak salt and water, upon the microscope

Fig. 9.— Muscular Fibres op Drone (Magnified 300 Times).
a, Part of Fibre, contracted ; b, Part relaxed ; tr, Trachea; n, Nerve.

stage, the wave of contraction may be seen play¬
ing along the fibre, almost as one observes it
in a garden worm, as it draws up the hinder part
of the body whilst moving onwards. The part
(as at b) quickly, by bringing together its plates,
assumes the appearance of «, while a extends itself,
plate by plate, until it is fully relaxed. The contrac¬
tion soon ceases, but I have watched it in operation
for at least two minutes. Nerves ( n , Fig. 9)

BEES AND BEE-KEEPING.

occasion this exceedingly beautiful rhythmic move¬
ment. Those desiring to study it, had better first
try the common gentle, its muscles not coming to
absolute rest till nearly half an hour after removal
from the body. Muscular fibres under a low power
of the microscope are easily recognised, on account of
their considerable size and striated (cross lined) ap¬
pearance. They are each covered by a remarkably
attenuated membrane, called sarcolemma, in which,
generally, a delicate tracheal tube takes its course.
Indeed, in the muscles of the wings every fibre has
its own particular tracheole (small trachea). The
muscular fibres, in this case, lie side by side, and are
arranged in bundles (fasciculi); across these pass air
tubes, parallelly arranged, which give off from their
sides these tracheoles at singularly regular intervals,
the latter being equal to the diameter of the fibres.
Each tracheole then follows the path of the fibre oppo¬
site to it with the uniformity of the rungs of a ladder.
This wonderful structure, like every other, could not
be properly examined without making us feel that
beauty in Nature is something more than skin deep.
Most muscles in the bee are attached direct to some
portion of the external skeleton, and, where distant
parts are thus to be connected by small muscles,
tendons are added, as we see in Fig. io, which re¬
presents the apparatus for opening and shutting the
jaw; here all the muscles have tendons, two of
which are exceedingly long. The striated fibres are
attached to the flattened terminal portion of the
latter, and are arranged in a plumose form, as seen
in the illustration.

GENERAL STRUCTURE.

How full of wonder and beauty is all this. A bee
runs into the hive, but it can only do so because
nerves stimulate, and a large number of muscles, each
containing many fibres, respond in accurate order, for
no joint of a single limb but moves as it ought in
obedience to the directing nerve-centres within the

Fig. 10.—Jaw of Queen, with Muscles (Magnified).

Ifm, Lesser Flexor Muscle; gfrn, Greater Flexor Muscle; lem. Lesser Extensor
Muscle; em, Extensor Muscle; m, Mandible; n. Notch; og, Gland
System (the Olfactory Gland of Wolff).

insect. How quickly movement follows upon move¬
ment, every step involving a complete circle of
changes. How tiny the muscles, how impalpable the
nerves, and yet large are they in comparison with
some others to be found in the same family of

44 -

bees AND BEE-KEEPING.

insects. The Anaphis, by example, possesses, like the
bee, its six legs, with nine joints each, its four wings,
and twelve jointed antennae, each supplied with its
proper muscles and nerves, and almost throughout
its structure part for part with its larger relative, and
yet its entire weight is less than xooooth of a grain.
What of its egg, carrying within its shell all the
directive essentials for evolving these pigmy marvels ?
The grandeur of the minute will as successfully hush
to silence the thoughtful man as the grandeur of
the vast.

CHAPTER IV.

NERVE SYSTEM.

Ganglionic Chain — Supra-cesophageal Ganglion—Rejlex
Action—Commissural Fibres—System of Larva and
Imago—Coalescence and Atrophy of Ganglia —
Cephalic Ganglia — Convolutions — Pedunculated
Bodies of Dujardin—Relation of Size of Brain
to Intelligence—Inferiority of Queens — Stomato-
Gastric and Sympathetic Systems — Variety of
Nerve Work.

The nerve system in insects (A, B, Fig. n),
whether in the larval or adult stage, consists mainly
of a series of rounded masses of brain-like substance,
arranged in the median line of the body, near to the
lower, or ventral side. These masses, called ganglia*
are united by two threads, seen in the figure, and
each of which is shown by the microscope to con¬
sist of a sheath, having. within it an immense number
of nerve fibres, serving to bring the separate ganglia
into union, by carrying impressions received by one
to all the rest. The front mass of all is not on the

Greek, yarfyKiov, a knot, or excrescence.

4 6

BEES AND BEE-KEEPING.

ventral side of the body, since the ganglion below
it (Fig. 12) sends off two short and curved straps
(really nerve bundles), called the oesophageal collar,
which embraces the oesophagus, or food passage, above
which the front mass, or brain, lies, denominated, in

Fig. 11.—A, Nerve System of Bee Larva ; B, Nerve System of Adult or
Imago (Magnified Five times); C, Ganglion (Magnified Sixty times).
a. Antenna ; mx, Maxilla; to, Mandible ; w, Wing ; 1, 2, 3, 4, 5, Ganglia; n, Nerves;
en. Nerves escaped from Sheath ; nl, Neurilemma ; gc, Ganglion Cell; cf, Com¬
missural Fibres ; rf, Reflex Fibres.

consequence of its position, the supra-oesophageal
ganglion. From reasons presently to engage our
attention, it is clear that this ganglion is the seat of
intelligence, and that impulses from it dominate the
rest, but that the latter are also capable, undirected,
of initiating properly concerted movements. A study

NERVE SYSTEM.

of this ganglionic chain gives us the key to many
facts, which must puzzle the apiarian who has not
become so overpowered by mystery that he has
ceased to inquire into the cause of anything.

Unfortunately, even in our humane system of bee¬
keeping, the tiny throng handled by such a Brobding-
nagian race as we relatively are, must now and again
meet with serious accidents, and we may have to
watch, with mingled astonishment and regret, the
rapid march of a headless victim, or the threatening
twisting of a detached abdomen, as its sting turns to
our finger, striving to execute the lex talionis. By look¬
ing to our illustration, we see that the decapitated one
is still possessed of much brain substance, distributed
through the body, and that the isolated abdomen
carries with it not less than five ganglia, brain portions,
so to speak, which initiate movements simulating
design, as we have seen. The loss of the head,
although it absolutely puts an end to whatever
amount of consciousness the insect possessed when
uninjured, reducing it thereby to the condition of a
machine, is not fatal, in a restricted sense, as we
have already hinted; and, curiously enough, drones in
confinement will sometimes live very much longer
without their heads than with them. After decapita¬
tion, when the irritation set up by the cut has sub¬
sided, they will remain perfectly quiet, but imme¬
diately they are disturbed they begin to move,
running about, and possibly attempting to fly. If
the body is turned over, a struggle is made to
adjust it, and a hair-pin, dipped in phenol or ammonia,
or any substance emitting an irritating vapour,

4 «

BEES AND BEE-KEEPING.

brought near to one side of the body, will imme¬
diately cause the residue of the insect to retreat in
the opposite direction. If the hair-pin be actually
put into contact with the body, the legs will at once
be set to work to rub from the part the cause of
annoyance. Facts like these are well known to
physiologists, and those who desire to extend their
acquaintance with them are referred to such works
as Dr. Carpenter’s Manual, or Huxley's Text-book.

At Fig. ii (C), we have an enlargement of one of these
ganglia, which shows it to be really double, one-half
belonging to the right, and the other the left, side of
the body. Let us suppose the lateral threads (n) (which
are turned upwards, for the sake of convenience, in
the illustration) to be provided to the right front leg.
If this member be pinched, touched, or influenced in
any way, an impression travels along the nerve until
the ganglion is reached, when the fibres take four
independent courses, all indicated in the figure—
some run forwards towards the head, others back¬
wards towards the relatively posterior nerve masses,
still others to the opposite half of the ganglion, thus
uniting in the impression the right and left sides
—such are called commissural fibres (cf Fig* i i) ;
and others, again (rf), after entering the mass of
the ganglion, and coming into contact with its
cells, return by the same side. The impression pro¬
duced immediately results in a movement, reflected
from the ganglion, without any intervention of the
action of the brain, and so such movements are called
reflex. The commissural fibres would originate action
on the opposite side of the body, while the fibres

NERVE SYSTEM.

running forward and backward would bring neigh¬
bouring ganglia, and possibly the brain also, into play.
Singular as this may appear to be, it is exceedingly
like that which is constantly occurring within our¬
selves. If, by example, an unfortunate soldier has a
shot wound dividing his spine, near to the middle of
its length, the whole of the lower part of his body
will be absolutely paralysed; he will be deprived of
both sensation and voluntary movements in his legs.
But if, now, his feet be tickled by a feather, although
he will feel nothing and know nothing of what is
occurring, unless informed by the eye, his legs will
plunge violently, and strive to remove the feet from
the source of titillation, because the irritation will be
carried by his leg nerves to those nerve cells of the
spine which are below the injury, and which exceed¬
ingly resemble the ganglia of the insect; and from
these impulses are reflected, resulting in energetic
action, with which his brain has, of course, nothing
to do, precisely as in the case of the decapitated
drone, which will by its legs forcibly push from it a
cause of annoyance, if such a word may be employed
in relation to that which has no consciousness. With¬
out going to so dread an example as a wounded
soldier, we may constantly trace in ourselves, or our
friends, movements which are purely reflex, resulting
neither from a sensation nor a mental impression, but
made, possibly, in the absence of both.

Nothing can be more striking than the difference
between the arrangement of the ganglia of the bee
larva and that of the same insect in the perfect
condition, unless we take into account the exceed-

50 BEES AND BEE-KEEPING.

ingly diverse circumstances of the life of the two,
the helpless dependence and quietude of the former
standing in marked contrast to the self-sacrificing
devotion and restless energy of that of the latter.
The changes, which fit the same nerve system for
such opposite conditions of being, are effected by
slowly-made modifications, commencing with the
creature's independent existence, but which are more
active and radical during the chrysalis, or pupal stage.

The business of the larva is to eat. It is produced
from an egg, which must be tiny, because the mother
laying it furnishes others in such prodigious number.
The minute body it possesses, when its first pap is
given to it, must increase in weight, as I have found
by careful experiment, about 1400 times during the
four days it feeds; and so its nerve system is now
principally distributed to its digestive apparatus and
to its spiracles. It claims, as yet, neither legs, wings,
nor eyes; nevertheless, during the period that the
one want of its lower existence is being met,
preparation is also being made for the higher endow¬
ments, new responsibilities, and enlarged enjoyments
of the future, for its nerve masses are already
coalescing, in order to become more perfect in their
functions ; they are concentrating their influence and
making the insect less vegetative. The larva has
seventeen embryonic ganglia (as seen in Fig. 13),
one supra-cesophageal, three sub-cesophageal—that is,
under the oesophagus, and also under the first-men¬
tioned ganglion, or brain—three thoracic, and ten
abdominal ; but, as it grows, and in an early stage,
the three sub-oesophageal and three last abdominal

NERVE SYSTEM.

in each case fuse into one, reducing the ganglia to
thirteen, as we have them in A, Fig. n, where, of
course, the two first ganglia, lying over one another
nearly, appear in actual contact. But when the old
digestive tube lining and contents have been cast
away (see page 22), and the chrysalis stage is reached,
metamorphoses proceed more rapidly. The two latter
thoracic and two first abdominal ganglia then unite,
forming a large and powerful nerve mass (as seen
at B, Fig. 11), still giving indications of its com¬
pound origin, and ‘ initiating the main external
activities of the insect by throwing out, from its
anterior parts, nerves to the second pair of legs
and the anterior wings, while its posterior half gives,
similarly, energy to the third pair of legs and the
posterior wings, the front legs receiving twigs from
the first thoracic ganglion. The original eight
abdominal ganglia, but now reduced to six (see
supra), suffer a further diminution in number by
the fourth and fifth melting together; and so we
find, in the worker, five abdominal ganglia, while, for
reasons given when treating of the queen and drone,
they have four each only. But let us not imagine
that coalescence and development are all that occur.
The larva had needs which it does not carry with
it when it leaves the cell, and so some structures
are atrophied, with the nerves supplied to them,
while their material, by absorption, is diverted to
other uses. These changes require much patient
investigation in order fully to trace them, but they
may in large part be easily seen by proceeding with
eggs and larvae of various ages, as stated on page 37.

BEES AND BEE-KEEPING.

We now pass to consider more in detail the
structure of the head, or cephalic ganglia, which
should show us the evident relation subsisting be¬
tween the wants of the animal and those curious
endowments which come to it, we know whence,
though we know not how, in the quietude and dark¬
ness of the little waxen cell.

Looking to Fig. n, B, we find in the head the
upper view of the supra-cesophageal, with the collar
uniting it to the sub-oesophageal ganglion, while, in
Fig. 12, we have a front and enlarged view of the
same. The former ganglion, or brain, is so soft and
transparent, that it is hardly possible to trace its form
without the use of some hardening agent, such as
alcohol, or chromic acid; but for a microscopic exami¬
nation of the character of its substance we must
operate upon a bee in a perfectly fresh state. The
upper part of the cranium being removed, we come
first upon salivary glands, numerous tracheae, and
tracheal sacs, covering up the brain, which is itself
inclosed in a double membrane, like the pia and dura
mater of higher animals; these stripped off, we reach
the pulpy material of the cerebral mass, consisting, for
the most part, of transparent globules, from -g-^Vg-th t°
5 p 0 0 th of an inch in diameter. If now we pour over
this some solidifying material—and, for popular work,
turpentine will answer well—we find it does not become
uniformly white and opaque, but convolutions, such as
seen at p, Fig. 12, begin to make their appearance
near to the ocelli, or simple eyes (0). By degrees,
removing the pulpy mass which covers over these
convolutions, we find the latter to be an interior sub-

NERVE SYSTEM.

stance, whiter and more solid, possibly corresponding
to the so-called white matter of the brains of verte¬
brate animals. The general form taken is seen from
B, which covers over, but still so as to allow to
appear, the so-called pedunculated bodies of Dujardin,
and is copied from an actual brain, compared with
the drawing given by that physiologist, in his admir¬
able Memoir* When these bodies are freed from
their surroundings, they are seen to bear a very

A B

Fig. 12.— The Brain, or Supra-cesophageal Ganglion.

A, Head, showing Brain, Ac., magnified ten times— o, o, o, Ocelli; a, a. An term*,
with Nerves ; p, p, Brain covering Pedunculated Bodies ; og, og, Optic Ganglion ;
s, Sub-cesophageal Ganglion. B, Supra-oesophageal Ganglion, or Brain,
deprived of External Membranes, magnified thirty times.

short peduncle, or stalk, pointing towards, and nearly
reaching, the median line; so that, although they do
not actually touch, they possibly bring the two
lateral halves of the brain into relation. These
stalks bear above them the convoluted lobes. A
granulous tubercle, placed in front of each of these
singular forms, and prominent in the ant as well as
the bee, is supposed by Dujardin to be especially

“Annales des Sciences Naturelles,” 3i»e S3rie, vol. xiv.

BEES AND BEE-KEEPING.

provided to receive the communications made by
these wonderful little creatures, by tappings on the
front of the head with the antennae.

Our figure shows that the brain sends three short
stalks to the ocelli (o), the centre one receiving its
nerves from the right and left side, while the brain
laterally passes into the two masses provided to the
large compound eyes ; on each side also, a well-
developed lobe, beneath, gives origin to a nerve
supplying the antenna, for Dujardin seems to have
been in error in supposing that these arose from
the pedunculated bodies.

Such, then, is the brain of the bee, declaring that
its owner is endowed, at least, with glimmerings of
intelligence. For, in those insects whose whole
course may be supposed to be simply instinctive,
the pedunculated body is not found ; in such, the
entire brain, and every ganglion, consists alone of
pulpy matter. Where the pedunculated bodies exist,
their bulk, as well as that of the antennae lobes,
seems to bear a direct proportion to the diversity of
action of which the creature possessing them is
capable. We have a progression in the size of these
appendages as well as in instinctive development in
passing— e.g., from the cockchafer {Melalontha vul¬
garis) to the cricket, on to the ichneumon, then to
the carpenter bee, and, finally, to the social hive
bee, where the pedunculated bodies form the -g-th
part of the volume of the cerebral mass, and the
-jj-foth of the volume of the entire creature, while, in
the cockchafer, they are less than the -^ t^ th part.
The size of the brain is also a gauge of intelligence.

NERVE SYSTEM.

In the worker bee, the brain is xr^th of the body ; in
the red ant, -g-g-g-th; the Melalontha, F5 - 0 - 0 th; the
Dytiscus beetle, 4 - 4 ^ Q th. And here a very curious
point arises. As we proceed, I shall have more
than once to point out a misconception, which would
appear to be all but universal amongst bee-keepers,
and to show that the queen is not superior to, but
greatly the inferior of, the worker; and the brain
bears evidence to this position, as that of the queen
is relatively small, as is also that of the drone.
The amazons, who support the political fabric of the
bee-hive, supply its food, bring up its young, furnish
its architecture, defend its property, administer
justice, and determine the how, when, and where of
new colonies, require greater endowments than the
males, and true female, who is largely aborted, so
as to be almost exclusively limited to the faculty of
reproduction.

Besides the principal nerve system of which we
have spoken—that of animal life—the bee, in common
with some other insects, is possessed of two other
systems, of less proportions, and more visible in the
larva than the adult, which give energy to the func¬
tions of organic or vegetative life. One is denomi¬
nated the stomato-gastric, and is provided with
numerous minute ganglia, which send nerve-fibres
into the organs of digestion, circulation, and respira¬
tion ; the other, corresponding probably to the sympa¬
thetic of higher animals, has, in each segment of the
body, a very small triangular ganglion, sending out
threads, which ultimately anastomose with those of
the previously considered abdominal chain.

BEES AND BEE-KEEPING.

It is exceedingly difficult, nay, rather, impossible,
although we are but thinking of the little bee, to
realise the wonderful complexity and capability of
this brain and ganglionic system, with its countless
nerve fibres and numerous nerve cells ever transact¬
ing the mystic and involved telegraphy of life,
receiving messages and transmitting replies with a
quickness as little to be conceived as that of the
electric current itself, besides stimulating and co¬
ordinating a great diversity of parts, and bringing all
into a conscious unit, and so endowing that unit, that
it is but part of a greater whole, which, in turn,
puts itself into true, determinate, and useful relation
to the world which forms its environment; but we
shall hereafter remember, that no muscle can move,
no heart throb, no organ of sense receive an im¬
pression, no gland secrete, and no digestion be per¬
formed, without the operation of some part of these
strange transparent threads, with their accompanying
ganglia.

CHAPTER V.

DIGESTIVE SYSTEM.

Need of a Digestive System — Food Solution —
Similarity of the Digestive Process in all Animals
— (Esophagus—Honey Sac — Stomach-Mouth—Chyle
Stomach—Gastric Teeth—Small and Large Intes¬
tines — Anal Glands — Embryogeny of Digestive
Tube — Structure and Use of Stomach-Mouth and
Chyle Stomach — Dufour’s Theory.

The primary object of a digestive system is to
supply the vitalising and formative material called
blood, which sustains in activity and builds up out of
its substance every tissue of the bodies of animals.
A little attention given to the process of blood
making in our own case, will well prepare the way
for a better realisation of the uses of the structures
we find in bees. Our food, whatever may be its
character, needs at first to be wholly or in part
brought into the condition of solution in water, and
only such part as is actually dissolved can be in
any way utilised ; eg., in ordinary bread, our typical
food, we have principally two substances, starch and
gluten, both of which may be soaked for any

BEES AND BEE-KEEPING.

period without dissolving, if decomposition be pre¬
vented ; but during the process of chewing or
mastication, glands, of which we, like our bees,
have three pairs, pour into the mouth, saliva, whose
principal office is to chemically change some parts
of our food, and notably starch, which, under its
action, begins to be formed into sugar, one of the
most soluble bodies furnished by the plant world.
After swallowing, the process of transformation goes
on, until at length all starch has disappeared, and
the sugar produced from it has, by absorption, got
into the blood current. This sugar, although derived
from starch, is still the representative of the honey
of the bee, while the gluten, the residue of our
bread, is the counterpart of her pollen—so similar
are our sources of sustenance. But gluten requires
a treatment distinct from that which the starch
received, for the former is not materially affected in
the mouth, but, passing into the stomach, the gastric
secretion acts upon it and so transforms it that a
new and soluble material, sometimes called albu-
minose, is produced, which can be, on account of its
liquid condition, transmitted to the blood, and that
mainly by the action of a multitude of minute
thread-like bodies, which cover the inner side of part
of the alimentary tube, and, so to speak, drink up
the dissolved, or, in other words, digested, nourish¬
ment. This glairy material, thin and transparent, is,
after absorption, carried up a narrow channel running
in front of the backbone, and poured at length into
a vein under the middle of the clavicle (collar-bone)
on the left side. Thus mixed with the blood, it is

DIGESTIVE SYSTEM.

quickly made into part of that fluid actually, and
now, with the rest, visits every muscle and organ
by the circulation, in order that it may nourish
and sustain. Far removed as we are from bees,
there still exists between us and them a most
helpful similarity of physical structure, and presently
we shall find that the salivary and gastric secre¬
tions perform precisely the same functions in both.

If the abdomen be pulled from the thorax of a
recently dead bee, until the integument, which is
really a part of the external chitinous envelope,
ruptures at the narrow junction of the two, called
the petiole or stalk, we shall almost uniformly drag
away with the abdomen a long thread-like form,
which is really the tubular oesophagus or gullet
( ce , Plate I.), running away from the tongue through
the head, neck, thorax, and petiole, about -g-in., until
it begins to enlarge within the abdomen.

It often happens that the rough surgical operation
just described will pull the digestive tube from the
abdomen, as well as expose the oesophagus ; but, if
not, but little skill is required in so opening the body
that the whole may be removed without much injury
to it.

Certain glandular products are added to the food
in the mouth, of which more hereafter; but the
oesophagus is only conductive in character, and is
narrow within the thorax, being -g^in. i n diameter.
The thorax, indeed, as the centre of locomotion, is
loaded with the strong muscles the legs, and espe¬
cially the wings, require, and so here no space can be
spared for the function of digestion. The enlarge-

BEES AND BEE-KEEPING.

ment just referred to {hs, Plate I.) is known as the
honey sac, and corresponds to the crop of most
insects. It is about -i-in. in depth and ^-in. in dia¬
meter when full of honey, of which it will hold a full
third of an ordinary drop. When nectar is gathered
by the foraging bees, it is simply held in store in this
cavity, the processes of digestion in no true sense begin¬
ning until the next chamber ( c.s )—the chyle stomach
—is reached. The bee having returned to the hive,
the cross muscles indicated in hs, Plate I., and LM,
TM, Fig. 14, by contraction, press upon the con¬
tained nectar and drive it back through the oeso¬
phagus into the cell of the comb, in the manner
described at page 18. But, if the mouth of the ox
that treads out the corn should not be muzzled, it is
clear that the little labourer should have an oppor¬
tunity of taking, of that it has gathered, for its
own support. To permit of this and much more, a
rounded body {p, Plate I.), of singular and beautiful
structure, about g^th of an inch in diameter, is placed
at the bottom of the honey sac. It can be easily
seen by the unaided eye, and is of pearly, yet
brownish, colour. This apparatus (which may be
more easily investigated in the Queen Bombi than the
hive bees, on account of its greater size in the former)
we shall carefully examine presently, at the moment
calling it the “ stomach-mouth,” a very appropriate
name, which Burmeister has given, and which suffi¬
ciently explains its use, for the bees’ food can be
taken through it at will, and as required* into the
chyle stomach. The latter bends much upon itself
in the worker, has a diameter of yL-in. and a length

DIGESTIVE SYSTEM.

6 :

of fin., but is straighter and smaller in the queen,
and its sides are, in all cases, banded with constric¬
tions that occur at regular intervals. The gastric
glands are placed in its walls, while the pollen grains
commonly found within it in abundance give to it a
yellow, or yellowish-brown, appearance. At its further
extremity it narrows considerably, and forms a pylorus
in passing into the small intestine (si, Plate I.), which
is here met by a considerable number of long and
narrow tubes (bt, Plate I.), lying in tangled spirals, but
which, nevertheless, enter the walls of the digestive
system with great regularity, their openings being
closely set, side by side, in a single encircling line.
They are, probably, excretory in function, removing,
like the liver in ourselves, impurities from the blood,
which are modified so as to be of service in the
work of digestion. Their structure is shown at
C, which gives a small portion of one of them,
magnified 450 times. These tubes are known as
Malpighian vessels, from the great anatomist, Mal¬
pighi, who discovered them; or are called biliary
or urinary tubes, according to the view which may
be taken of their office. The intestinal lining mem¬
brane here undergoes an interesting modification; it
is arranged in a number of longitudinal ridges, and
is set with small, though hard, chitinous teeth (D,
gt, Plate I.), frequently double-pointed, each about
~ 2 '60 o in - * n length. They are most easily seen in a
newly-hatched bee, before any food has been taken
to interfere with the view. The object of these,
in my opinion, is to abrade the growing points of
any pollen grains which have not sufficiently yielded

BEES AND BEE-KEEPING.

to the action of the chyle stomach, so that their
nutritious contents may be duly appropriated. To
enable them to accomplish this, a strong coat of
ring muscles is provided, thus equipping the bee
with a rudimentary gizzard, beyond which the small
intestine lies, somewhat twisted, and not quite
uniformly placed in different bees; its diameter,
which varies little throughout its length, is about
s^in. It is shown in cross section in D, Fig. 14.
The muscles supplying it are remarkable, and may
be conveniently studied under the microscope after
staining with eosin, or even ordinary red ink. The
colour of its contents is perceptibly darker than
that of the chyle stomach, while the pollen
grains, which in the former are but little altered,
are here generally damaged in the cellulose cover
(see page 10), and are frequently broken up com¬
pletely, as they have had to pass the mill of the
gastric teeth. The small bowel suddenly, and
afterwards more gradually, expands into the colon,
or large intestine {li, Plate I.), which is often
swollen and dark in appearance, because its trans¬
parent and colourless sides show clearly its contents,
which have here that disagreeable odour too well
known to bee keepers who have given liberty to
bees that have endured the confinement and worry of
a long journey. At the commencement of the colon
are placed six longitudinal, brownish, fleshy plates
(/, Plate I.), which appear to be both valvular and
glandular in action ; they are protuberant on the
inner side, and are formed by an invagination of the
intestinal walls, the whole of the layers of which

DIGESTIVE SYSTEM.

take part in their structure. Tracheae densely
ramify in these tubercles, and a large nerve is
supplied to them. M. Leydig compares these fleshy
plates to the tracheal lamellae forming the rectal
branchiae, or gills, of the aquatic larvae of the
libellulidae, or May flies.

Embryogeny (the science of the development of
embryos) shows that, whilst the bodies of insects
are being fashioned within the egg, the digestive
tube is formed in three parts, as follow: The
mouth and anal extremities by invaginations (see
page 34) of the external skin, the central portion
from a modification of the yolk sac, which is met
at its ends by the continued deepening of the
posterior and anterior invagination. The parts have
now no communication, their ends being blind,
but are placed like two half sausage skins, with
a whole skin in the centre. The separating walls
at length undergo absorption, and the tube be¬
comes single and united, passing through the body
from end to end ; the annexed organs are then
marvellously added to this simple tube, which forms
gradually, at definite spots upon its wall, prolonga¬
tions, at first like the fingers of a glove, but by
degrees assuming the involved structure they possess
in the adult insect. In this manner within the
bee egg the spinning glands (Fig. 15), which
subsequently become System 3 of the salivary
glands (Fig. 16), are formed at the anterior part,
and at the posterior, similarly, the Malpighian, or
urinary tubes. An arrested development, however,
in bees, hornets, and wasps, causes the middle

6 4

BEES AND BEE-KEEPING.

bowel to remain blind at the posterior extremity,
which may be well seen at Fig. 13, representing the
developing larva within the egg membrane; here the
anterior invagination (fb) has already made junction
with the middle bowel {mb), but the after bowel {ab)
remains separate, and Will continue to do so until
the commencement of the chrysalis condition, so that
the larva, usually so prolific of dejections, in the case
of the above-mentioned insects passes nothing. If
it were otherwise ordered with bees, by example, the
embarrassment would be great where the larva lies

Fig. 13.— Bee Larva before Hatching (Magnified Forty times).

eh, Chorion, or Egg Skin ; ga, Ganglia; s.ga, Supra-oesophageal Ganglion; jm, Jaw
Muscles forming; c, Nerve Collar; fb. Fore Bowel; mb, Middle Bowel;
ab, After Bowel.

in a cell surrounded on all sides by liquid food ;
besides which, the honey we now so much value
would be made unacceptable, through possible con¬
tamination from an uncleanly nursery. I have already
explained my discoveries respecting the way in
which the accumulated residua are at length got
rid of so as to leave the larva unsullied (page
22). These surprising changes, humbling us by
showing us how little we know, and how much there
is to learn, may, without difficulty, be witnessed by
those possessing a stock of bees to furnish eggs, and

DIGESTIVE SYSTEM.

a microscope to examine them. Let us now investi¬

gate in detail the stomach-mouth and chyle stomach.

Fig. 14.—Honey Sac Stopper, or Stomach-Mouth, &c. (Magnified Fifty times).

A, Front View of One of the Four Leaflets of Stopper, or Stomach-Mouth—
l, Lip-like Point, covered by Down-turned Bristles (b ); sra, Side Membrane.
B, Longitudinal Section of Stomach-Mouth, with Continuations into Entrance
of Chyle Stomach— l, l, Lip-like Ends of Leaflets; *, Setae; Im, Longitudinal
Muscles; tm, Transverse Muscles, in Cross Section; cl, Cell Layer of Honey
Sac; LM and TM, Longitudinal and Transverse Muscles of same; nc.
Nucleated Cells of Tubular Extension of Stomach-Mouth into Chyle
Stomach; lm' and tm', Longitudinal and Transverse Muscles of Chyle
Stomach ; c, c, Cells covered within by an Intima. C, Cross Section of
Stomach-Mouth— m, Cross Section of Muscles seen at lm, in B; tm, Trans¬
verse Muscles surrounding Stomach-Mouth. D, Cross Section through Small
Intestine —a and m. Longitudinal and Surrounding Muscles.

We have already learnt that the first of these
enables the bee to store honey, which, although

BEES AND BEE-KEEPING.

carried within her body, does not enter her digestive
system, and that by the means of it food can be
taken into the chyle stomach as required, p, Plate
I., shows us the stomach-mouth, as seen through
the transparent walls of the honey sac, its form being
not unlike an unopened flower-bud with four sepals.
If it be carefully removed from a recently-killed bee,
and examined by a simple lens, its lips or leaflets
may frequently be observed opening and shutting
with a rapid snapping movement. A more remarkable
object than this under a low power of the microscope
can scarcely be imagined. The oesophagus, honey-
sac, and chyle stomach, should be removed together,
and placed on a glass slip, the microscope stage being
made horizontal. No cover glass should be used, but
sufficient very weak salt and water added. The whole
object will exhibit, for at least fifteen minutes, muscu¬
lar contractions of a most instructive kind, while the
gaping and snapping of the stomach-mouth, and the
passing onwards of food, is often noticed. If the bee
operated upon has just previously been fed with
honey stained with some aniline dye, the effect is
enhanced. By closing the oesophagus I have fre¬
quently succeeded in getting, not only food, but even
bubbles of air, gulped down into the chyle stomach,
and, by carefully pressing upon the stomach-
mouth with the side of a needle, the lips may be
forced open, and food passed on into the stomach
beyond. One leaflet being separated from the rest,
we find it strongly chitinous within, and fringed along
its margin (A, Fig. 14) by downward-pointing,
fine, but strong bristles. At B we have the longi-

DIGESTIVE SYSTEM.

tudinal section of the whole apparatus, with its

entrance into the chyle stomach, somewhat as figured
by Schiemenz.* It is provided with two sets of
strong muscles, one ( Im ) running perpendicularly
along the backs of the leaflets, and, by their con¬
traction, pulling asunder the lips, and permitting a
passage of food from the honey sac to the chyle
stomach; another {tin), in cross section in the
figure, running round the whole, and perfectly

closing it at the will of the bee. The figure in

like manner shows the two muscular layers (LM,
TM) of the honey sac, by the united contraction
of which the gathered nectar is driven out into

the cells of the comb for general consumption.
C, as- it gives the form in horizontal section, with
the opening and closing muscles, makes somewhat
clearer the beautiful mechanism of the stomach-
mouth, the utility of which is so conspicuous : for the
bee can eat whenever and wherever she likes : when
she departs from the old home, with all its stores pro¬
vided against a “ rainy day,” and commits herself, with
her companions, to the vicissitudes which the swarm
must encounter, she can minimise her risks by carry¬
ing, in the honey sac, sufficient food for a week's
necessities, either using it rapidly in the production
of wax, or eking it out, should the elements prove
unfavourable for the gathering of new supplies ; and
in winter, when departure from the cluster is im¬
possible, she can, at infrequent intervals, as oppor¬
tunity arises, so charge herself from the honey cells

* See “ Uber das Herkommen des Futtersaftes und die Speicheldriisen
der Biene .”—“ Zeitschrift fur Wissenschaftliche Zoologie,” Bund 38.

BEES AND BEE-KEEPING.

that her wants will always be supplied, and her ability
to produce heat be uninterrupted.

But, besides these beautiful adaptations, another use
has been suggested. Leon Dufour taught that the
larvae of bees are nourished by an ejection into their
cells of semi-digested food from the chyle stomach of
the nurses, and this idea, unsupported as it is by
evidence, has gained all but universal acceptance.
Schonfeld* explained the stomach-mouth in con¬
formity with this opinion, but recent investigations
have more than ever convinced me of the erroneous
nature of Dufour’s theory. Schonfeld at first alto¬
gether failed to observe that the stomach-mouth is
prolonged into the chyle stomach by a tube contain¬
ing a layer of nucleated cells ( nc , B, Fig. 14),
beyond which extends an extremely delicate mem¬
brane (intima), which Schiemenz is confident can
have no other object than to prevent the return of
digesting matters into the honey sac, his opinion
being that, except when food is passing through it,
this tube must collapse completely, being pressed on
one side, and flattened. But microscopic examination
and experiment have shown me that, although the
tube of intima interferes with regurgitation, as Schon¬
feld is forced to admit, still it may float in the
stomach, and preserve its cylindrical form notwith¬
standing pressure, so that its presence rather makes
regurgitation improbable than impossible. Schonfeld
has also left unnoticed the down-pointing bristles
(/, A and B, Fig. 14), which would, by straining,

* See translation in British Bee 'journal, 1st Tulv, Kth SeDt.
15th Oct., and 1st Dec., 1883. 0 v '

DIGESTIVE SYSTEM.

effectually prevent the passing upwards of any solid
particles, such as pollen grains, whole or broken,
even could the difficulties previously mentioned be
overcome.

But it will be seen that these explanations are
partly negative, giving us no reason for the presence
of either tube extension or down-pointing bristles, since
a mere sphincter (or ring of closing muscles) would,
by contraction and relaxation, have either totally pre¬
vented regurgitation, or permitted it, if necessary, and
also have enabled the bee to take such food from
the honey sac as it might at the time being contain.
It is clear, then, that either parts have been added
which are not requisite, or that some function exists
which has hitherto escaped observation. Surely it is
the latter. Dissecting bees from the hive, young and
old, ordinary nurses and queen grub feeders, starved
and fully fed, gave me no help in this matter, beyond
showing the extraordinary complexity and variety of
movement of which the stomach-mouth is capable;
but those that were engaged in gathering yielded the
solution.

On the Compositse, as well as many other orders,
bees suck up nectar, in conjunction with much
pollen, and, examining the honey sac of one work¬
ing upon a single dahlia, e.g., the outside wrinkled
membrane (sm, A, Fig. 14) is seen to continually run
up in folds, and gather itself over the top of the
stomach-mouth, bringing with it, by the aid of its
setae, the large pollen grains the nectar contains.
The lips (/, /, B, Fig. 14), now opening, take in this
pollen, which is driven forwards, into the cavity made

7 o

BEES AND BEE-KEEPING.

between the separating lips, by an inflow of the fluid
surrounding the granules. The lips in turn close, but
the down-pointing bristles are thrown outwards from
the face of the leaflet, in this way revealing their
special function, as the pollen is prevented from re¬
ceding while the nectar passes back into the honey
sac, strained through between the bristles aforesaid,
the last parts escaping by the loop-like openings
seen in the corners of C. Fig. 14. The Whole pro¬
cess is immediately and very rapidly repeated, so
that the pollen collects, and the honey is cleared.
Three purposes, in addition to those previously enu¬
merated, are thus subserved by this wondrous me¬
chanism. First, the bee can either eat or drink from
the mixed diet she carries, gulping down the pollen
in pellets, or swallowing the nectar, as her necessities
demand. Second, when the collected pollen is driven
forwards into the chyle stomach, the tube extension,
whose necessity now becomes apparent, prevents the
pellets forming into plug-like masses just below p,
Plate I., for, by the action of the tube, these pellets are
delivered into the midst of the fluids of the stomach,
to be at once broken up and subjected to the
digestive process. And third, while the little gatherer
is flying from flower to flower, her stomach-mouth is
busy in separating pollen from nectar, so that the
latter may be less liable to fermentation, and better
suited to winter consumption. She, in fact, carrier
with her, and at once puts into operation, the most
ancient and yet the most perfect and beautiful of all
“honey strainers.”

The chyle stomach is lined by an intima, or inner

DIGESTIVE SYSTEM.

7 1

membrane, carrying a cell-layer ( c ), the cells com¬
posing which appear to be of two kinds, having
distinct functions, one secreting a digestive fluid
(gastric juice) from the surrounding blood into the
stomach, so that the contents of the pollen grains
may be made fit for assimilation, by a transformation
not unlike that liquefying gluten in our own case ;
the other absorbing the nutrition as prepared, and
giving it up into the blood—these cells representing
the absorbent vessels of ourselves and the higher
animals generally. Outside this cell-layer comes a
propria, or outer membrane, and, beyond this, two
muscular coats, one {tin') of ring muscles, the other
(Im ) of longitudinal muscles, which, by their appro
priate contractions, originated by the stomato-gastric
nerve system, churn the contained food, and move it
onwards past the, several constrictions previously men¬
tioned, and which are commonly twenty-three in
number, until the pyloric extremity is reached. The
process of absorption continues in the intestines till
only waste products and indigestible matters remain,
and these are ejected by a muscular action, which
can only be effectively employed, in the case of the
worker and drone, when the insect is on the wing.
The queen presents an exception to this rule, which
will, hereafter, require an explanation.

The view here suggested, that the brood is not
nourished by regurgitated material, l£ads at once to
the question, How, then, is it fed ? No satisfactory
answer can be given until we study the gland struc¬
tures.

CHAPTER VI.

SALIVARY (?) GLANDS OF BEES.

Discovery of Glands—Plans for Dissection—Position
and Connection of Systems No. /, No. 2, No. 3 —
Details of the Systems—Intracellular and Inter¬
cellular Glands—System No. 4 of Schiemenz —
Uses of the Glands — Variations in Queens—Brood
Food — Brood Weaning — Royal Jelly — Queen
Ra ising — Fertile Wo rkers — Egg Production — A
New Theory Respecting Queen Food — Queen
Dejections — How Queens are Stimulated into
Laying by the Nurses—System No. 1 a Chyle
Gland -— Retarded Laying Explained — Feeding-
Groove.

In 1811, Ramdohr announced the discovery of a pair
of salivary glands in the thorax of bees, whilst two
other pairs were found by H. Meckel, in 1846; and
yet dense ignorance respecting them is common to
the present day, even such an accomplished German
apiculturist as Berlepsch failing to mention them,
while Cook only in his last edition (1884) calls
attention to the existence of two pairs, which he
tells us were “first discovered by Mr. Justin Spald¬
ing/’ It is not a wonder that the rank and file of

GLAND STRUCTURES.

bee-keepers are as much in the dark as those to
whom they look for leading. Leydig* and Sieboldf
did much to elucidate the structure of these glands,
but their methods of dissection were not sufficiently
refined to enable them to properly locate them in
the body of the bee; Siebold, in particular, falling
into serious mistakes on more points than one,
followed by Girard, who does not appear to have
himself made any dissections.

It has already been stated that the larva secretes
its cocoon from a gland, which reappears, in a

A, Gland— p, Propria; cl, Cell Layer; i, Intima; r, Reservoir; sc, Secreting
Cells. B, Transverse Section of Reservoir. C, Transverse Section of Secreting
Layer, with Interior Lumen, or Cavity.

modified form, in the adult. This gland is seen
at Fig. 15. Its product, as is usual with insects,
remains perfectly liquid so long as it is stored in the
reservoir (r), but quickly hardens after it is drawn
out into threads, although not so rapidly but that the
several filaments where they cross each other partly
fuse together, and so much strengthen the gossamer
blind which the larva elaborates (see Fig. 4). The
secretion itself is derived from the blood by the
action of the cells (sc), seen in the cross section C,

* “ Archiv fur Anatomie Physiologie,” &c., Leipzig, 1859.
t “ Mittheilungen iiber die Speichelorgane der Biene,” 1872.

BEES AND BEE-KEEPING.

and surrounding a small tube, or lumen. These cells
have, of course, an absorbing surface on the outside,
while their proximate faces secrete a liquid silk,
which, as formed, is passed into the reservoir, of
which B is a cross section, where it collects in con¬
siderable quantity before the time of spinning.

By inserting a needle into the mouth of a worker
bee, and passing it upwards, behind the front wall
of the head, the latter may be so opened that its
salivary (?) glands, in a partly broken condition, may
be obtained for examination ; but if the attachments
and entire forms are to be investigated, we must
proceed as follows: Partly fill some shallow recep¬
tacle, such as a pomatum-pot, or large pill-box, with
melted bees’, or paraffin, wax. When cold, with a
hot wire melt a little bath in the centre of the
waxen surface, and then insert the bee we wish to
dissect, so placing in this case that one side of the
head is submerged. By a second application of the
wire, re-melt the wax in the, neighbourhood of the
head, using no more heat than is necessary to secure
thorough adhesion, and now cover with water or
glycerine. A powerful light and a good watch¬
maker’s eye-glass (secured round the operator’s head
with a tape, when it can be pushed up on to the
forehead if not required) will permit of reason¬
ably good dissection, although, of course, better
results can be reached by using a Stephenson’s
erecting-binocular-microscope — the instrument with
which all the dissections for this work have been
made. The bee thus securely held by the wax, both
hands are free to manipulate. Now, with a needle-

GLAND STRUCTURES.

knife (made by heating a large needle, beating it

Fig. 16.—Salivary (?) Glands of the Bee.

A, Salivary (?) Glands, Systems No. 1, No. 2, and No. 3, magnified fifteen times— sv.
Salivary Valve (of Systems No. 2 and No. 3) at Root of Tongue; Ip, labial
Palpus ; mx, Maxilla ; so, Salivary Opening of System No. 1 in Hypo-pharyngeal
Plate; no, Openings in Plate for Termination of Taste Nerves; ce, (Esophagus ;
sd, Salivary Duct; b, Junction of Ducts of System No. 2 ; c, Junction of Ducts
of System No. 3; sc, sc, Salivary Sacs; fl, Front Lobe; bl, Back Lobe ; a, Chitinous
Duct, with Spiral Thread. B, Single Acinus of System No. 1, magnified seventy
times— n, Cell Nucleus; st, Salivary Track ; d. Large Duct. C, Single Pouch, or
Acinus, from System No. 2— a, Propria, or Outer Membrane ; sc, Secreting
Cells. D, Termination of System No. 3—1 2, 3 4, lines marking Termination of
Section ; d. Duct, in section ; sc, Secreting Cell, in section ; n, Nucleus.

flat, and afterwards sharpening upon a hone, and

7 6

BEES AND BEE-KEEPING.

inserting into a wooden handle), cut carefully round
the compound eye, and lift it off. Curiously folded,
and passing round the optic ganglion, we have a
long whitish body, which a facetious friend com¬
pared to ropes of onions. It is one side of the
System No. i of Siebold (Fig. 16). Behind this,
and extending from the top of the head downwards,
we find packed inimitably a second gland system
(No. 2), consisting of many pouches, joined by canals
to a common duct, which may be followed until it is
discovered to enter another duct (6, Fig. 16) running
backwards and forwards in the body. Tracing this
channel towards the thorax, we see it enter the neck,
and immediately after bifurcate or fork (c, Fig. 16).
Following the line of one of the two ducts, we come
upon a reservoir (sc), leading backwards to another
gland system (No. 3), of singular structure, with
two lobes, lying in the front of the thorax on each
side of the body. The position of all these systems
is well seen in Plate I. The operation here described
is not likely to be accomplished with one bee, and I
spent many days, and spoilt many specimens, before
getting the glands in their entirety, with their con¬
nections ; but I have good reason for supposing that
these successful dissections are. unique. Leaving out
of view for the present a fourth gland, attached to
the jaw (Fig. 10), and which Siebold failed to note,
let us proceed to examine in detail the systems to
which he gave name.

Taking pains to secure an entire right or left gland
of System No. 1, we find it to consist of an inelastic,
transparent, central tube or duct, without branches,

GLAND STRUCTURES.

and of the uniform diameter of from -g-^in. to ^^ in.,
surrounded through its length, which is fully once
and a half that of the entire body of the bee, by
between 1000 and 1200 berry-shaped bodies, called
acini, of which one is much enlarged, B, Fig. 16.
In these acini the secretion is produced by cells, which
develop, perform their function, and pass away, to
be succeeded by others. The cells forming each acinus
are surrounded by a bag-like membrane, or propria,
through which the blood passes continually, to supply
the material out of which the secretion is elaborated.

System No. 1 is intracellular in type— i.e., every
part of the surface of each cell is absorbent, so that
the secretion it furnishes has to be removed from its
interior by a duct, which enters its wall, becomes sur¬
prisingly delicate, and takes within a lengthened,
sinuous course, bringing itself in contact with the
cell plasma. These chitinous tubes, each about
Tgwo'n. in diameter, after leaving the cells, pass
parallelly through an enveloping tube ( st , B), towards
d, where, by independent perforations, they enter
the main duct, which at this point raises itself into a
sort of papilla, having a sieve-like end. In the red
ant, a similar gland (K, Plate VII.) has its cells free,
the propria being wanting. Its form as given should
be studied. Tracing this duct onwards towards the
mouth, we find it enter a pouch, or ampulla, lying
at the side of, and beneath, a plate which forms
what may be termed the mouth-floor (Fig. 17).
The part {pi) of this plate drops as a flap towards,
and joins, the upper extension of the tongue, so
that food passing over the latter can be uninter-

7 8

BEES AND BEE-KEEPING.

ruptedly carried back, over the hypo - pharyngeal
plate, to the oesophagus, or swallow beyond. Ex¬
tremely strong and dark-coloured horn-like forms
run backwards, and converge on each side, while
near their ends are seen prominences which give
attachment to the protractor pharyngis muscle (pp),
which, by shortening, throws forward the whole ar¬
rangement, bringing the front of the plate close tip

Fig. 17.—Hypo-pharyngeal Plate (Magnified Twenty-five times).

A, Plate in Worker— pi, Plate Leaflet; No. 1, Main Duct of System No. 1, with
Acini removed; ptn, Papillse of Taste Nerve ; oa, Opening of Ampulla for
Escape of Secretion; pp, Protractor Pharyngis Muscle; p, Pollen Grains in
Commencement of (Esophagus. B and C, Halves of Pharyngeal Plate from
Queen and Drone (lettering as before.)

behind the epipharynx ( g, Plate II.). We find what
at first might be taken for apertures at tn, but a
careful examination shows these to be delicate papillae
of the taste nerve, which runs beneath the pharynx,
and passes its terminating fibrils into them. But
our main point now is the discharge opening ( oa )
of System No. I, a portion of whose duct, with its
continuation through the plate, is represented, though

GLAND STRUCTURES.

deprived of its acini, which are broken away by the
least violence.

Systems No. 2 and No. 3 are intercellular, like the
spinning gland, whence they are derived— i.e., the cells
are arranged around, a cavity, towards which they pre¬
sent their secreting surfaces, while they absorb material
from the blood by that portion lying next the propria
[a, C, Fig. 16). The secretion passes forwards in a
manner made obvious by the illustration. When the
several ducts begin to unite, they develop an interior
spiral thread like to that of the tracheae, both in
purpose (page 34) and appearance. Their presence
led Fischer to suppose these glands to be lungs. In
System No. 3 these threads are especially strong, as
indicated at a A, d D. They pour their contents into
a sac (sc, A), curiously covered hy star-shaped plates.
The ducts of both systems uniting, as previously
described, form a single channel, passing onwards
through the mentum (mt, Fig. 18), or chin, into the
tongue, where it terminates in a salivary valve (sv, A,
Fig. 18), from which the saliva is pumped out during
the action of sucking, an operation which may be
artificially performed after the death of the bee.

System No. 4 of Schiemenz, or the olfactory gland
(Riechschleimdriise) of Wolff * (og, Fig. 10), closely
resembles No. 1 in its minute structure, being intra¬
cellular, and, in consequence, very active. It has its
aperture immediately within the mandible, is singu¬
larly large in the queen, smaller in the worker, and
still less in the drone.

* “Nova Acta Acedemia Naturae Curiosorum,” vol. xxxviii., “Das
Riechorgan der Biene,” Dr. O. J. B. Wolff.

BEES AND BEE-KEEPING.

A question of surpassing interest, but immense
difficulty, now presents itself, viz., What is the purpose
served by each of these glands? Admitting, for argu¬
ment’s sake, that the view taken in the last chapter,
of the office of the stomach-mouth, is correct, we have
three or four distinct functions to be performed by
such structures as we are now considering. First, a
secretion to assist digestion; second, to change the
cane sugar of the nectar of flowers into the grape
sugar of honey, and possibly also convert starch into
sugar (both of these functions are performed by one
salivary secretion in our own case) ; third, to soften
and make plastic the wax plates formed on the under
side of the abdomen, so that they may be elaborated
into comb, and also possibly serve as a vehicle in the
moulding of propolis, or the application of it as a
varnish ; and, fourth, the production of a brood food.

Without dogmatising, my investigations into this
question lead me somewhat confidently to point to
System No. i as actually having the latter office.
For it is first worthy of remark, that this gland
—the largest and most active—is only found in the
worker bees. By referring to Fig. 17, we note that
B, the hypo-pharyngeal plate of the hive queen, has
scarcely any perforation, and that the merest trace
of duct is attached to it, having clearly no secreting
power. It is peculiarly important, as well as
interesting, to observe here, in a parenthesis, that,
the higher the quality of the queen, the further will
she be removed from the worker in this matter,
poor queens, hurriedly raised, really possessing this
gland in an extremely rudimentary form, while those

GLAND STRUCTURES.

with the largest ovaries have even the plate im¬
perforate, while no trace of duct is discoverable, just
as in the case of the drone plate (C). But taking
a queen Bombus (page 13), engaged in establishing
a nest, when she does feed her own brood, we find
this particular gland strongly developed; whilst in
other bees, such as the mother Megachile centun-
cularis (page 9), which secretes no wax, but raises
her own young, we still see it, though of smaller
form. We thus get some evidence that rendering
wax plastic is not the duty of this gland, but that
the feeding of brood is. Again, examining a young
worker employed in nursing, we find this gland
turgid, and in the highest state of activity; while in
the old bees of a broodless stock it is much
shrunken, at the same time that glands No. 2 and
No. 4 retain their normal size. Coupling this fact
with the larger dimensions of No. 2* and No. 4 in
the queen, and remembering her need of assimilation
in order that her eggs may be produced, we shall not
be far wrong in ascribing to No. 2 and No. 4 a diges¬
tive function. In other words, they are truly salivary
in character, which position is further supported by the
existence of these glands in less development in the
drone. But to return ; microscopical examination of
the food given to very young larvae reveals no trace of
a pollen grain, and shows that it resembles in nothing
any part of the contents of the chyle stonlach of the
nurses. It is, on the contrary, just such a fluid as a

* Siebold, followed by Girard, says that No. 2 is small in the queen;
but this is clearly an error. In many scores of queens dissected, I have
uniformly found it larger than in the worker, and often containing sacules.

BEES AND BEE-KEEPING.

secretion might be. As, however, the larva gains size
and power, the process of weaning commences, and
its food undergoes a change, having now undoubted
pollen, honey, and water added to it—the glandular
secretion being, of course, gradually withdrawn. The
pollen grains, moreover, are living, and are generally
found in a growing condition, proving that they have
never entered the stomach of the nurse, and, certainly,
that they are not semi-digested, and so utterly con¬
tradicting the Dufour theory. In the case of the queen
larva, I discover that weaning is not adopted, but that
secretion, commonly, though, . as I hold, erroneously,
called royal jelly, is added unstintingly to the end; so
that, at the close of the feeding period, an abundance
of highly nutritious food, which I apprehend does not
intrinsically differ from that at first given to the
worker larva, remains, and to which the chrysalis for
some time adheres, possibly continuing to draw from
it, by osmose (fluid diffusion), material which aids
its development. The queen larva does get a very
small addition of pollen, the residue of which collects
in the middle bowel; but this seems to be rather
accidental than otherwise.

The first brood food, or pap (page 19)—I am almost
tempted to say bee milk—is, then, a highly nitrogenous
tissue-former, derived from pollen by digestion, and has,
apparently, a singular power in developing the genera¬
tive faculty; for I find drone larvae receive much more
of it than those of workers, to whom any accidental
excess possibly gives the power of ovipositing, as we
find it in the abnormal fertile worker. From these
considerations, I have been led towards a theory, the

GLAND STRUCTURES.

evidence in favour of which has accumulated until I
cannot but regard it as established. It is, that the
queen, if not always, at least during the time of egg-
laying, is fed by the workers from the secretion of
gland No. 1, with possible additions from some of
the others.

It has been already stated (page 26), that the
queen, at certain periods, has the power of producing
between 2,000 and 3,000 eggs daily. Each one of
these is -^in. long, -^j-in. in diameter ; and a careful
calculation shows that 90,000 would occupy a cubic
inch, and weigh 270 grains. So that a good queen,
for days, or even weeks, in succession, would deposit,
every twenty-four hours, between six and nine grains
of highly developed and extremely rich tissue-forming
matter. Taking the lowest estimate, she then yields
the incredible quantity of twice* her own weight
daily, or, more accurately, four times, since at this
period more than half her weight consists of eggs. Is
not the reader ready to exclaim, What enormous
powers of digestion she must possess, and, since pollen
is the only tissue-forming food of bees, what pellets
of this she must constantly keep swallowing, and
how large must be the amount of her dejections!
But what are the facts ? Dissection reveals that
her chyle stomach is smaller than that of the
worker, and that, at the time of her highest efforts,
often scarcely a pollen grain is discoverable within it,
its contents consisting of a transparent mass, micro-

* Queens vary considerably in weight, small ones, in the winter, not
exceeding 1$ grains, while a few, in the middle of the spring-laying, will
turn the scale at 3 grains—feeding adding fully half a grain more.

H 2

BEES AND BEE-KEEPING.

scopically indistinguishable from the so-called royal
jelly; while the most practised bee-men say they never
saw the queen pass any dejections at all. These con¬
tradictions are utterly inexplicable, except upon the
theory I propound and advocate. She does pass de¬
jections, for I have witnessed the fact; but these are
extremely watery, and are voided with great energy,
while she rests, back downwards, on the bottom of the
comb. At least,.this has been her position when I have
noted the occurrence. Moreover, although her stomach
is small, her urinary tubes are exceedingly active and
large, adding further confirmation to my position, as
these enable her to rid herself of the great excess of
water a secretion diet would supply. We thus see
that her digestive function is performed by proxy, the
residuary matter of the pollen required to produce
her eggs being, under this exceedingly beautiful ar¬
rangement, carried from the hive in the bodies of
the feeding bees, to be expelled in mid-air—so
wondrous are the devices by which the require¬
ments of these creatures are met.

When first hatched, the queen is not noticed—she
is but one of the multitude, requiring nothing special,
and has, of course, no feeding attendants, but takes
her nourishment from the cells, like the rest, and
empties her bowels when on the wing, like a worker.
Her weight does not at this time increase, or she would
become incapable of all the soaring of the marital
trip. Now her stomach always contains pollen ; but,
from the hour of her impregnation, she is the subject
of watchful attention, the younger bees gathering
about her, not to form her body-guard, as writers

GLAND STRUCTURES.

have generally fancied, but to minister to her
necessities ; and her weight from this time rises very
rapidly, her ovaries developing under the influence
of what I shall call chyle* food, which, two or three
days after impregnation, her stomach contains in
quantity, while all trace of pollen has disappeared;
but if I be not correct, this is the period above all
others when large quantities of pollen should be
undergoing digestion. I have sacrificed many queens
just when at their very highest value, for the purpose
of settling, as far as may be, this important inquiry,
with results most uniform and confirmatory. Here,
too, I imagine we get the key to the retarded
laying, always noticed when a queen fails in im¬
pregnation ; i t is because the bees t hemselves fail
in administering that kind of nourishment which
stimulates the ovaries. And, in addition, we learn
how it is that the colony have under control the
laying powers of their queen, stimulating her or not,
as circumstances warrant.

It is necessary now to observe that honey, like
sugar, is what the physiologist denominates a “ force-
former," and, as such, is needed by the q'nfeen
to supply her activities, and so queens may be seen
to dip their heads into honey cells and there drink ;
while a queen not laying may be supported for some
time upon sugar syrup alone.

The ducts of Systems No. 2 and No. 3 are so placed

* System No. 1 Siebold has unfortunately called “salivary,” in
ignorance of the facts to which I now call attention. Since this term
is very misleading, I shall refer to this gland hereafter as the chyle
gland.

BEES AND BEE-KEEPING.

that their secretions are given up only as the tongue
is protracted or extended as for sucking (see Figs.
18 and 19), while the peculiarity of the position of
the discharge opening of the chyle gland (No. 1) is
just such as my theory requires. There exists upon
the worker's tongue, and upon the worker’s only
(Plate II., and gr B, Plate III.), a feeding-groove, or
narrow trough, on to which honey is brought by the
compression of the honey-sac when one bee feeds
another. Just at the back of this feeding-groove, when
the tongue is retracted (see A, Fig. 19), lies the plate
into which the chyle gland (No. 1) opens; and, by
a combination of most extraordinarily complicated
muscles, between thirty and forty in number, the chyle
can then be taken from this feeding-gland, and placed
upon the groove at once, for the benefit of the queen;
or it can also, when the tongue is doubled back in
repose, be brought into the right position for feeding
the larvae from the mouth, as the latter lie at the
bottom of the cells. There are yet other considera¬
tions pointing in the same direction, which must
not be anticipated until we come to practical matters.

Since I am not conscious of a single fact which
appears in any way to-throw doubt upon what I have
advanced, while every point examined has brought
to it additional corroboration, I leave the argu¬
ment to speak for itself, lest this chapter become
unduly lengthened. Every advanced bee-keeper will
see its practical importance, and that it dispels the
shades of many mysteries; while the naturalist will,
in delight, realize that his bee is more a wonder of
wonders than he has before imagined.

CHAPTER VII.

TONGUE AND MOUTH PARTS.

Endo-Skeleton of Thorax and Head — Meso-cephalic
Pillars—Mouth of Bees — Mandibulee, Labrurn , and
Labium — Mentum not Tubular — Labial Palpi
and Maxillx—How Large Quantities of Honey
are Taken—One Use of the Epipharynx—The True
Sucking Tube of Bees—Sucking Small Quantities
of Nectar — Sheath — Rod—Centre and Side Ducts
of Tongue — Bouton — Pouch—Solidity of Tongue:
How Simply Determined — Necessity of Pseudo-
Tubular Form—How to Distend Pouch Artificially —
Queen and Drone Tongue—How Tongue is Folded
out of View—Folding by the A ndrenidae—Nectar
Converted into Honey — Why Bees Take Thick
Syrup Slowly—Feeding Brood in Cells—Reasons
for Wedge Shape of Head.

As we now commence another section of our anato¬
mical studies, leaving the internal organs for those
that, in large part, appear at the surface, some intro¬
ductory remarks are necessary.

The skeleton of insects, although external, is not
exclusively so— e.g., in the thorax of bees, subject as

BEES AND BEE-KEEPING.

it is to the strain of the leg and wing movements,
corrugations and plaitings, supplemented by internal
webs [see Plate I.), are provided, to give the needed
rigidity, just as the engineer secures the same by
corrugating his sheet iron, and adding webs to his
girders. Beside these, between the meso and meta¬
thorax, lies a stiff extension—really a plate bone—
called the meso-phragma, to give solid attachment to
part of the muscles of the organs of flight.

Fig. 18.—Longitudinal Section Through Head, just Outside Right
Antenna (Magnified Fourteen times).

a, Antenna, with Three Muscles attached to mcp, Meso-cephalic Pillar; cl,
Clypeus ; Ibr, Labrum, or Upper Lip; No. 1, Chyle Gland (System No. 1 of
Siebold); this Gland really runs in front of the Meso-cephalic Pillars, but here
the latter are kept in view ; o, Opening of same; oc, Ocellus, or Simple Eye;
eg, Cephalic Ganglion ; n, Neck; th, Thorax; ce, (Esophagus ; sd 2, 3, Common
Salivary Ducts of Systems No. 2 and No. 3; sd 2, and sd 3, Salivary Ducts of
Systems No. 2 and No. 3 respectively; sv. Salivary Valve ; c, Cardo; ph, Pharynx ;
lb, Labium, or Lower Lip, with its Parts Separated for Display ; mt, Mentum,
or Chin; mx, Maxilla ; Ip, Ip, Labial Palpi; l, ligula; b, Bouton.

In the head, which has to sustain the heavy pull of
the jaw and tongue muscles, besides defending the
brain and delicate glands, corrugation is prevented
by the presence of the very large compound eyes,
with their essential regularity of outline, an element
of weakness, unless some device were introduced to

TONGUE AND MOUTH PARTS.

prevent it; and so an endo-skeleton, or inner frame¬
work, is added, its most important part being a
pair of strong rods—the meso-cephalic pillars (mcp,
Fig. 18)—running from front to back, and attached by
their extremities, just outside and below the antennae,
and at the rear of the head, beneath and on each
side of the occipital opening, the orifice through
which junction is made with the thorax. In a lower
position are placed a second pair of internal girder¬
like forms, supplying attachment to the cardos ( c,
Fig. 18), which, in turn, act as levers to the lower lip,
of which we shall speak presently.

Let us, whilst carefully consulting our illustrations,
direct our attention to the well-worn but ever fresh, the
charming but still difficult, problem of the mouth parts
of the bee, about which, perhaps, more has been written,
and more error propagated, than any other organism
of equal size in this wondrous creation. This verit¬
able crux of anatomists, with its delicate complexity,
was far beyond the powers of the older instruments
of research, and so we regretfully leave the achieve¬
ments of bygone worthies, with all their testimony to
patience and to conscientious adherence to truth,
notwithstanding many contradictions, since we now
cover all their ground, and far more. Recently, the
brilliant monograph of Dr. O. J. B. Wolff* has added
much to our knowledge, while investigations by Mr.
Chambers and Mr. J. D. Hyattt have given new
light on the other side of the Atlantic. From the
two latter, Professor Cook, in his “ Manual of the

* See Footnote, page 79.

f American Quarterly Microscopical Journal, vol. i., page 287.

BEES AND BEE-KEEPING.

Apiary/’ derives his chief facts; but, unfortunately,
he has added to these several statements which are
astoundingly inaccurate, and from which our minds
must be freed at once, if we are to have any intelli¬
gent idea at all of the tongue of the bee. Since his
book has been largely read in this country, and its
teaching generally accepted as the result of those
microscopic examinations of which it continually
speaks, I must refer to these and other errors, in the
interest of naturalists and bee-keepers, as we pro¬
gress ; and, happily, they are not those which, on
account of their difficulty, leave room for diversity
of opinion, but such as can easily be made clear,
even, in some cases, without a microscope.

The mouths of all insects have the jaws moving
sideways. The caterpillar, which carves our cabbage-
leaves with an industry which cannot secure our ap¬
proval, places itself at the edge of the leaf, driving one
jaw through the upper, the other through the lower sur¬
face thereof. In like manner, our bee has its mandi-
bulae, or outer jaws ( m , Plate II.) at the right and left
of the upper lip, or labrum, which depends between,
and is provided with a row of delicate feeling hairs.
These jaws are notched, in the queen and drone
(Plate IV.), as we find them in wild bees, but are
entire (i.e., not notched) in the worker (Plate II.),
are very powerful, and serve, amidst many purposes,
to be noticed in due course, for biting, as well as
thinning out wax shreds in comb building. Beneath
the upper lip appears (at g) the front of the epi-
pharynx, covered by a delicate white membrane, and
containing an abundance of nerves, endowing it with

TONGUE AND MOUTH PARTS.

some special sense, which Wolff concludes to be
smell—an opinion in which I cannot agree, for
reasons given hereafter. The under side of the
mouth-opening (mo, Fig. 18) is formed by the labium,
or under lip (lb), which is seen to embrace a number
of parts, carried by its basal portion, or mentum (ml.
Fig. 18, or B, Plate III.). The mentum lies beneath the
head, and somewhat behind it, and can, within con¬
siderable limits, be moved backwards and forwards.
It is strongly chitinous below, is articulated to the
head by means of the sub-mentum, or lora (lo, Fig. 18,
or /, B, Plate III.), and contains the muscles (a, a, rb,
B, Plate III.), which can draw the tongue, or ligula,
partly back into it, and also conveys the salivary duct
(sd, Fig. 18), which opens by a valve (sw) at the base
of the ligula. Otherwise the mentum has no opening,
is filled with blood, has nothing to do with the
oesophagus, or swallow, and is, of course, not tubular,
as Cook states. The ligula is not a continuation of
the mentum in front, but has its roots within the
latter, from, which it is withdrawn by the action of
a muscle (the protractor linguae), when the tongue
is outstretched for sucking. Attached to the mentum
at its front margin, and possessing at this point a

hinge joint, we have on each side of the ligula a

labial palpus (Ip, Fig. 18 and Plate II.). It consists

of four joints, the two upper being large, the two
lower very small, and provided with elaborately

contrived feeling hairs. Outside these we find the
maxillae (mx, Fig. 18, and Plates II. and III.), attached
to the sub-mentum (the right maxilla has been
removed in Fig. 18), and having a chitinous portion

BEES,AND BEE-KEEPING.

hollowed out, and fitting against the side of the
mentum, which it partially embraces with very stiff
hairs, so that they hold it in position. The maxillae
have feelers or palpi ( mxp , Plate II.), which appear
in the hive bee to be aborted and functionless,
although in the Andrenidae they are usually well-
developed. Not far below the feeler, the maxilla,
like the labial palpus, has a hinge, separating the
higher, tougher part (the stipe) from the lower,
more delicate, and transparent portion forming the
lacinia, or blade. The sections C, D, E (Plate III.)
show that the maxillae and labial palpi normally em¬
brace the tongue before and behind respectively, so
that together they may form a tube, within which
the tongue is placed. If we now remember that the
tongue can be drawn back in part into the mentum,
while the embracing parts cannot, we see at once
that the tongue has the ability to move up and down
within the formed tube; and again, that as the maxillae
are attached to the lora, the maxillae may move back¬
wards and forwards upon the labial palpi.

With the outline before us of the main parts of
this complex structure, let us endeavour to under¬
stand the methods of its action, and the purposes
to which it has been adapted. First, How are large
quantities of honey taken ? The ligula, when ex¬
amined by a low power, is found to be covered by
a sheath [sh, B and G, Plate HI.), densely clothed
with hairs, regularly arranged, in the worker, in from
90 to 100 transverse rows, of which the queen and
drone only possess from sixty to sixty-five. These
hairs, most symmetrically placed, and passing through

TONGUE AND MOUTH PARTS.

a beautiful gradation of form, from row to row, are
short and triangular in shape near the base of the
organ, long and spiny about the middle, smaller
and more flexible near the apex, while amongst them
are found hairs with a bulbous structure, provided with
a nerve which constitutes them touch organs, or true
tactile hairs (th, I, Plate III.). The high elasticity
of the ligula, depending upon the structure of a rod
running through its centre, allows it to be used as a
lapping tongue when any considerable quantities of
syrupy food are at command. As it then sweeps
backwards and forwards, the front side turned down,
the gathering hairs (gh, Plate III.) get loaded, while
the labial palpi and maxillae are so placed round it
as to form a perfectly airtight tube. C, D, E,
Plate III., show the palpi with the hairs crossed
behind, while the hyaline plates ( hp , C) of the two

maxillae lie over one another. Each maxilla is

beautifully furnished with a line of hairs ( h ) in front
of it, and a groove at its back ; both of these act as
mechanical stops, and accurately adjust the position
of the two maxillae, preventing them, as they are

drawn together by their proper muscles, from ap¬
proaching too nearly whichever plate may happen to
take the front position. But although the tube so
made up is airtight, it cannot act as a suction pipe,
because it is open above, as may be seen by

reference to Plate II. But now the front extension
of the epipharynx (g) closes down to the maxillae,
fitting exactly into the space they leave uncovered,
and thus the tube is completed from their termination
to the oesophagus.

BEES AND BEE-KEEPING.

An important question now arises : How is this act
of sucking performed? Some have supposed that the
suction originated in the mouth ; while Cook calls
the honey-sac the “sucking stomach,” using an old,
but extremely misleading, title, for this wrinkled
membrane could no more exert suction than could a
balloon extract gas from the main. Comparing D
with E, Plate III., which are cross sections at the
same point, we see that the space (through which
the nectar must travel) surrounding the ligula, and
between the palpi and maxillae, is three times as
great in the former as the latter. This greater space
is obtained by arching the maxillae above the ligula,
and so causing the former to retreat from the palpi.
Here, then, is the origin of the sucking. The tube
is made to expand rhythmically above, the nectar
follows up into the space thus provided, and then, as
this contracts again, travels on into the pharynx, as
B, Fig. 19, will make clear.

Second, How is nectar taken when smaller quantities
only are obtainable ? A more minute investigation of
the ligula is now essential.

The so-called sheath of the tongue is highly chiti-
nised, stout, and very elastic, while the hairs which
clothe it are broad at their bases (gh, I, Plate III.),
and pre-eminently suited to gather up syrupy fluids
by capillarity, but utterly unfit for collecting minute
quantities of nectar, only reachable, perhaps, by the
extremity of the tongue, as would be the case in
most flowers, especially those with a tubular corolla.
Here, then, another and surprisingly beautiful con¬
trivance meets us. Taking a cross section through

TONGUE AND MOUTH PARTS.

the middle of tongue (G, Plate III.), we note, first,
that the strong sheath, sh (from which the gathering
hairs have been removed, for the sake of clearness),
passes round the tongue to the back, where its edges
do not meet, but are continuous with a very thin
plaited membrane (pm), covered with minute hairs.
This membrane, after passing towards the sides of
the tongue, returns to the angle of the nucleus, or
rod, over the under surface of which it is probably
continued. The rod passes through the tongue from
end to end, gradually tapering towards its extremity,
and is best studied in the queen, where I trace many
nerve threads and cells. It is undoubtedly endued
with voluntary movement, and must be partly mus¬
cular, although I have failed completely in getting
any evidence of striation. The rod on the under
side has a gutter, or trough-like hollow (cd, the
central duct), which is formed into a pseudo-tube
(false tube) by intercrossing of back hairs. It will
also be seen that, by the posterior meeting of the
sheath, the space between the folded membrane (sd)
becomes two pseudo-tubes of larger size, which I
shall call the side ducts.

These central and side ducts run down to that
part of the tongue where the spoon, or bouton ( 6 ,
Plate II. ; K, Plate III.), is placed. This is provided
with very delicate split hairs (E, Fig. 24), capable of
brushing up the most minute quantity of nectar,
which, by capillarity, is at once transferred by the
gathering hairs (which are here numerous, long, and
thin) to two side groove-like forms at the back of
the bouton, and which are really the opened out

BEES AND BEE-KEEPING.

extremity of the centre and side ducts, assuming,
immediately above the bouton, the form seen in F,
Plate III. The central duct, which is only from
■g^p-in. to To V o 'i n - diameter, because of its smaller
size, and so greater capillary attraction, receives the
nectar, if insufficient in quantity to fill the side ducts.
But good honey-yielding plants would bring both
centre and side ducts into requisition. The nectar is
sucked up until it reaches the paraglossae ( pa , B,
Plate III.), which are plate-like in front, but mem¬
branous extensions, like small aprons, behind ; and
by these the nectar reaches the front of the tongue,
to be swallowed as before described. Thus, then,
the bee is equipped to take advantage of all sources
of supply. She can gulp down big draughts, or sip
a stream of nectar so fine that 600 miles of it will,
when evaporated, store but a ilb. section box.

We are now, then, in a position to settle the ques¬
tion that has disturbed the minds of entomological
and apicultural writers for the two centuries and
more elapsing since the time of Swammerdam—Is
the bee’s tongue solid, or is it tubular? The problem
has been one of the highest difficulty to the micro¬
scope, depending upon the determination of the
nature of the back of the central duct, and authori¬
ties have been pretty equally balanced respecting it.
I agreed entirely with Wolff, that the duct was a
trough, and not a tube; but this left the question
one of authority or observation, and so still open to
debate. But, luckily, a form of experiment occurred
to me which settles the dispute most conclusively,
and in such a way that a 10s. microscope will answer

TONGUE AND MOUTH PARTS.

as well as a better one. Bees have the power, by
driving blood into the tongue, of forcing the rod out
from the sheath, and distending the wrinkled mem¬
brane, so that in section it appears as at H, Plate III.,
the membrane assuming the form of a pouch, given in
full-length at A. It will be seen at once that this
disposition of parts abolishes the side ducts, but
brings the central duct to the external surface. The
object of this curious capability on the part of the bee
is, in my opinion, to permit of cleaning away any
pollen grains, or other impediment that may collect in
the side ducts. The membrane is greasy in nature,
and substances or fluids can be removed from it as
easily as water from polished metal. If, now, the side
of a needle, previously dipped into clove oil in which
rosanilin (magenta) has been dissolved, so as to stain
it strongly red, be touched on the centre of the rod,
the oil immediately enters, and passes rapidly upwards
and downwards, filling the trough. I could not resist
laughing, while my pulse certainly went faster, as I
realised the absurdly simple means which put that
matter within the grasp of the tyro, with his simple
lens, which had kept many learned doctors wrangling,
notwithstanding all their appliances. It is a pseudo¬
tube, then, and nothing more. Hairs cover it, and
these permit of its being entered by the side.

The impossibility of cleaning, and so the tre¬
mendous risk, or, rather, the certainty, of clogging,
which a closed tube would involve, does not appear
to have disquieted those who have been wrongfully
describing the bee’s tongue as tubular; but there is
yet another consideration, which should show us the

9 8

BEES AND BEE-KEEPING.

surpassingly beautiful suitability of the form now ex¬
plained, for, although a tube would be fatal, a tubular
form is essential, as we shall presently see. Large
quantities of nectar may safely enough be gathered
outside the ligula in contact with the air, but small
quantities so collected would, in warm and dry
weather, inevitably thicken into a glue, which would
at once fix the poor little tongue, and for the time,
if not altogether, stop its labours. Sparing supplies
are, therefore, made to pass through the centre and
side ducts. The former is doubly protected from
evaporation, and by its channel the tiniest stream
would be as limpid at the mouth as when it left the
bouton ; while the side ducts, although more exposed,
only permit of such inconsiderable evaporation that
no risk whatever exists, while the pollen grains that
may perchance enter can be cleaned from it in the
manner we have already seen.

Cook says the sac to which we just now directed
our attention “ may be distended with nectar, as it
has connection with the tube of the mentum ”—

statements utterly at variance with the anatomy of

the parts, and capable of complete refutation by a
very elementary experiment. If hatching brood be
removed from the hive to a position too dry or too
cool, some of the bees will only succeed in freeing
their hea.ds from the cells, and will, in this position,

die. Of course, they have not fed, and yet in the

majority of cases the tongue pouch will be fully dis¬
tended, because the enforced stillness of the body,
and activity of the head, has determined an excess of
blood into the latter. Many will like to repeat my

TONGUE AND MOUTH PARTS.

experiment upon the central duct, for which stained
glycerine or honey may be used instead of oil,
although the latter is to be preferred. Let me ex¬
plain, then, how distended pouches may be obtained
without sacrificing any brood. I reflected that blood
might artificially be driven into the pouch ; and, taking
a recently dead worker, pinched the thorax between
thumb and finger from behind forwards; instantly
the pouch filled out, returning into position as the
pressure was relaxed. In nine cases out of ten, this
experiment with workers succeeds; with queens it is
difficult, on account of the extreme hardness of the
thoracic plates.

The queen's tongue is not only short, but the cen¬
tral and side ducts are not drawn out to the delicate
terminations we find in the worker, exactly as we
should have expected, since she has not to lap
minute quantities of nectar from the bottoms of blos¬
som cups, but simply to take food from cells, or,
more commonly, from the tongues of her attendants.
As is also clearly, necessary, the feeling hairs in
her case are far more developed than in that of
the worker, enabling her to determine in the dark¬
ness of the hive the exact point of the feeding bee's
body that she is approaching. The drone's tongue,
in like manner, is short, but is not highly sensitive.

All observant bee-keepers must have noticed that
the long and lithe tongue of their little assistants
disappears in a most astonishing fashion when no
longer required. Fig. ig will make clear the method
of its folding. At B we see it extended. It is
retracted by partly withdrawing it into the mentum,

100

BEES AND BEE-KEEPING.

as before stated, and carrying the mentum itself
backwards, by which movement the delicate skin
lying between the two secretory openings bends upon
itself, and the tongue, embraced by the maxillae,
doubles back behind the head, as at A. So that, in-

Fig. 19.—Ideal Sections Through Tongue (Magnified Twelve times).

A, Tongue Fully Retracted. B, Tongue Outstretched for Sucking; lettering as
Fig. 18. C, Ideal Line of Pharynx and Tongue in Activity. D, Method
amongst the Apidce, or Long-Tongued Bees, of Folding Tongue in Repose—a,
Articulation at Base; b, Bouton, or Point. E, Method amongst the Andre-
nidce, or Short-Tongued Bees, of Folding Tongue in Repose; lettering as D.

stead of presenting one sweep line (C), it is divided
into three between a and b (D). The whole family
of the Apidae thus turn the point of the tongue
backwards, while the Andrenidae (see page 8) turn it
forwards by a single doubling.

A most beautiful adaptation here becomes evident.
Nectar gathered from blossoms needs conversion into
honey. Its cane sugar* must be changed into grape
sugar, and this is accomplished by the admixture of

See “The Chemistry of the Hive,” Otto Hehner, F.C.S., British Bee
yournal, 15th Nov., 1883.

TONGUE AND MOUTH PARTS.

IOX

the salivary secretions of Systems No. 2 and No. 3,
either one or both. The tongue is drawn into the
mentum by the shortening of the retractor linguae
muscle, which, as it contracts, diminishes the space
above the salivary valve, and so pumps out the
saliva, which mixes with the nectar as it rises, by
methods we now understand.

Bees, it has often been observed, feed on thick
syrup slowly; the reason is simple. The thick syrup
will not pass readily through minute passages with¬
out thinning by a fluid. This fluid is saliva, which
is demanded in larger quantities than the poor bees
can supply. They are able, however, to yield it in
surprising volume, which also explains how it is that
these little marvels can so well clean themselves from
the sticky body honey. The saliva is to them both
soap and water, and the tongue and surrounding
parts, after any amount of daubing, will soon shine
with the lustre of a mirror.

The tongue is kept fully drawn back during the
feeding of brood, and the salivary valve is now not
only closed, but shut completely out of action by the
folded skin (A, Fig. 19) ; while the chyle gland
(No. 1) is brought up close to the tongue root, and
into the precise position for feeding from between the
mandibles. The wedged shape the tongue and head
take together is highly suggestive when the form of
the cell at the bottom of which the larva lies is
remembered. This wondrous tongue has no speech,
but yet who so dull that cannot hear its thrilling
little voice, speaking as unmistakably as the stars
discourse the language of the immensities?

CHAPTER VIII.

ORGANS OF SPECIAL SENSE—ANTENNAH AND EYES.

Difficulty of Subject — Touch, Taste, and Sight —
Antennae—Movements of Scape and Flagellum —
Feeling Hairs—Smell Hollows—Special Formation
in Queen—Conoid Hairs—Six Distinct Structures
—Hearing Organs—Sir J. Lubbock’s Experiments
—Proofs of Bees Hearing—Smelling — Experi¬
ments with Male Moths—Comparison of the Sexes
—Number of Smell Organs—The Equipment of
the Drone—Compound Eye — Pigment—Hexagonal
Facets — Methods of Examination — Crystalline
Cone—Nerve Elements of Eye—Mosaic Vision —
Microscopic Experiment — Stemmata—Colour Sense
—Albino and Eyeless Drones—Eyes of the Sexes
Compared — Conclusion.

The study of the special senses of creatures so far
removed from ourselves as bees cannot but present
great difficulties, quite apart from the minuteness of
the structures involved; for it is by no means impos¬
sible—nay, it is, rather, highly probable—they possess
modifications of sensibility which we can no more
truly realise than can the blind imagine the difference
between red and green.

ORGANS OF SPECIAL SENSE.

103

We have already seen that bees are not wanting
in the sense of touch, although it does not reside in
the exo-skeleton, but in multitudes of tactile hairs,
distributed as required. The sense of taste, too, is
possessed by the mouth and tongue, the hypo-pharyngeal
plate in the first being pierced for nervous exten¬
sions (page 78), while the second has, on each side, at
its root, thirty-two papillae, which are entered by
nerve end cells, just at the spot in which the nectar
meets the salivary secretion. In addition, analogy
seems to point to the nerve endings of the epipharynx
as also being taste organs. As the use of the eye
is obvious enough—upon the supposition that bees
may enjoy the senses common to animals higher in
the scale of creation—we have yet to look for organs
of hearing and smelling, and it will be well for us
to bear this in mind as we investigate the antennae,
commencing with those of the worker.

These cylindrical organs ( a, Plate II.) are inserted
near to each other, just above the margin of the
clypeus, and consist of two main portions—a single long
joint, denominated the scape, and eleven succeeding
short joints, called the flagellum. By a hemispherical
cup, the scape is articulated to the cranium, the latter
being moulded into a concavity (shown by shading in
the Plate), to permit of the widest range of motion on
the part of the former. The movements of the scape
are controlled by three muscles, seen lying behind the
antenna root (Fig. 18). One throws it outwards, the
second raises and draws inwards, the third depresses.
Two muscles in the scape itself ( Im, dm, Fig. 20)
move the flagellum. The second, third, and fourth

104

BEES AND BEE-KEEPING.

joints are well clothed with hairs, but otherwise are
dissimilar to one another, and to the remaining eight,
these last having a common structure of a highly
complicated character. The joints are not telescopic,
but are articulated (as at B, Fig. 20), with a central
opening between each, whose width is rather less
than half of their total diameter. They have only

very slight relative movement, and this is brought
about by contractile connections (a, B). If the eight
joints referred to be examined by the microscope, it
will be seen at once that the front and back faces are
totally unlike. The back is sparsely covered by
regularly placed hairs, somewhat curved, and pointing
downwards, whilst here and there hairs larger and
quite straight are seen (c 1 , B, Fig. 22). The front
is similarly furnished, but the spaces between the
hairs are here filled with oval discs, depressed in the

ORGANS OF SPECIAL SENSE.

105

centre, and having two or three faintly visible con¬
centric rings. These structures, although the first to
strike the eye, are the most difficult to understand,
and so had better be considered last.

Schiemenz* has examined the antenna by sections,
and since my own work shows me the accuracy, beauty
and success of his, I adopt his drawing (Fig. 21), with
only one or two modifications. The smaller hairs
iff) are loosely set into the chitine framework of the
antenna by a delicate ring, into which rises a nerve
end cell with a distinct nucleus. These hairs, standing
above the general surface, constitute the antennae
marvellous touch organs ; and, as they are distributed
all round each joint, the worker bee in a blossom cup,
or with its head thrust into a cell in the darkness of
the hive, is, by their means, as able accurately to
determine as though she saw; while the queen,
whose antenna is made after the same model, can
perfectly distinguish the condition of every part of
the cell into which her head may be thrust. The
last joint, which is flattened on one side, near the
end, is more thickly studded, and here the hairs are
uniformly bent towards the axis of the whole organ.
No one could have watched bees without discover¬
ing that, by the antennae, intercommunication is ac¬
complished ; but for this purpose front and side
hairs alone are required; and the drone, unlike the
queen and worker, very suggestively, has no others,
since the condition of the cells is no part of his
care, if only the larder be well furnished. The

* See Footnote, page 67.

BEES AND BEE-KEEPING.

106

conoid hairs ( c Fig. 21) are probably only highly
specialised feeling bristles, and are found in greatest
number at the extremity of the antennae. Each
antenna carries no less than six distinct structures,
viz., two forms of hairs not sensory on the scape and
upper joints, the ordinary sensory hair, the conoid
bristle, the elliptic discs, and, lastly, another structure
about to be described, and which is extremely likely

/, Feeling Hair ; s. Smelling Organ ; ho, Hollow ; c, Conoid Hair ; hi, Hypodermal
Layer ; n, n, Nerves, in Bundles; ar, Articulation ; c', Conoid Hair (Magnified
800 times).

to escape attention. Its external appearance is given
{ho, B, Fig. 22), and consists, superficially, of
minute holes, from -g-j^-g-in. to i^oo 0 ^ n - across, each
surrounded by a bright reddish ring. The micro-
scopist had better choose the antenna of a young
drone, and use a -|in. or - 1 -in. objective, with a Lieber-
kiihn, and then—unless as patient as microscopists
proverbially are—he will be as likely to lose his temper
as find the object. It is situated at the lower part
and outer side of the last six or seven joints of the
flagellum, but is found in greater abundance as we
get towards the end, the terminal joint carrying a

ORGANS OF SPECIAL SENSE.

107

patch of perhaps twenty, which creep round towards
the front. We have two of these in section {ho, Fig.
21), where we see the hole leads into a large cavity,
beyond which extends a widening cone. I am not
convinced that this cone is filled, as Schiemenz
supposes, for I regard it as an organ of hearing, its
larger size in the drone, with his possible need of
distinguishing the sound of the queen’s wings, and
its position on the outer sides and ends of the
antennae, seeming to me to favour this opinion. It
also appears to answer to parts considered to be
auditory organs in other insects.*

Sir J. Lubbock has commonly been regarded as
asserting the total deafness of bees; but, in a corre¬
spondence of some years since, the distinguished
investigator assured me his position was negative, as
he merely failed to get evidence of bees hearing. Sir J.
Lubbock’s experiments I cannot but regard as most
inconclusive, since tuning-forks, whistles, and violins,
emit no sounds to which any instinct of these crea¬
tures could respond. Should some alien being watch
humanity during a thunderstorm, he might quite simi¬
larly decide that thunder was to us inaudible. Clap
might follow clap without securing any external sign
of recognition ; yet let a little child with tiny voice
but shriek for help, and all would at once be awakened
to activity. So with the bee : sounds appealing to its
instincts meet with immediate response, while others
evoke no wasted emotion. In practical matters, the
hearing of bees is not only often obvious, but must

* “ Ants, Wasps, and Bees,” page 227, Sir J. Lubbock (Inter¬
national Science Series).

108 BEES AND BEE-KEEPING.

be taken into account— e.g., when a swarm is about to
be transferred to its permanent abode from its tem¬
porary one, many will stick to the sides of the latter,
after the bulk have been thrown out, and these, by their
buzz, will distract those that are running in at the new
hive door. The removal of the stragglers to a distance
will end the disturbance, which will be renewed if
they be returned to their former position. Some
years since I was present in a tent where an expert
had driven (see “Driving") five or six stocks, and
nearly a pint of lost bees had collected for mutual
comfort on a piece of damp canvas, at the bottom of
the tent pole, against which the last skep was made
to lean, as it was stood, quite late in the evening, on
a table for operation. No sooner did the bees in this
skep set up the well-known roar, than those on the
canvas, so still hitherto, faced upwards, unhesitatingly
ascending the pole, and settling on the outside of the
roof of the receiving skep. This circumstance I
remember as affording, to all who witnessed it, con¬
clusive evidence of hearing.

In the progress of the present we moderns have,
perhaps, too confidently condemned all the past.
The conflict of the key and warming-pan of old
swarming days has called forth some good-humoured,
but possibly not always philosophical, banter, for I
confess I think, that in its day, it had its value.
Piping of queens, whatever be its cause, seems to
point to a sense of hearing, for it appears to be a
sound made for an object, and not the result of some
necessary movement. Whether the organs we have just
considered be those of hearing or not, the possession

ORGANS OF SPECIAL SENSE.

109

of this sense by bees, of which much evidence will
subsequently come before us, cannot be doubted.

We have now to consider the “smell hollows” (in
cross section, s, Fig. 21), covered by a thin layer lying
over a goblet-formed cavity beneath,, into which passes
a nerve end cell, clearly unlike that provided to the
feeling hair (f). These oval forms are distinctly not
tactile, on account of their depressed position, but, for
reasons now following, almost certainly olfactory.
That the sense of smell is possessed by the antennse
simple observations would appear to favour. If bees
have food presented to them, the ends of the antennae
are, in alternation, brought close to it before the tongue
is advanced. If it contains even a small quantity of
an objectionable body which evaporates, the bee imme¬
diately retreats; but if the added substance be non-
vaporisable, such as corrosive sublimate, the antennae,
although used, do not detect its presence. The tongue,
however, immediately suffers, of which evidence is
given by the hurried departure, and the earnest efforts
made to clean away the cause of offence.

About three years since, near Bagshot, I carried
across a heathy plain, skirted on each side by firs, a
fresh female Emperor moth, lying at the bottom of a
muslin bag. After travelling about a mile, I retraced
my steps; and although, during several days’ hunting
for wild bees in the same locality, I had not seen an
" Emperor,” the males now met me, constantly flying
fearlessly up to the muslin bag—my companion, a
collector, having a very busy time. Similar facts every
naturalist could relate. The antenna; of male moths
are exceedingly large and extended in surface, and

BEES AND BEE-KEEPING.

the evidence that these are marvellously sensitive
to some emanation from the female is universally
accepted. But what of our bees ? Let us compare the
antennae of the sexes. The flagellum, which is the
sensory part of the antenna, I find, by careful measure¬
ment of many individuals, to be, on an average, in the
queen, y^-g-in. in diameter, yj-in. long; worker, yyyin.
diameter, -|-in. long; drone, -gyin. diameter, -±-in. long.
So that the sensory surfaces in the three cases are very
nearly in the ratio—queen i, worker 2, drone 3. Yet
the male, as his habits would lead us to suppose, has
only about 2000 feeling hairs, being the one-sixth or
one-eighth of the number of those possessed by the
worker. But what of the smell hollows? In the case
of the worker, the eight active joints have an average
of fifteen rows of twenty smell hollows each, or 2400
on each antenna. The queen has a less number,
giving about 1600 on each antenna. If these organs
are olfactory, we see the reason. The worker’s neces¬
sity to smell nectar explains all. We, perhaps, exclaim
—Can it be that these little threads we call antennae
can thus carry thousands of organs, each requiring its
own nerve end ? But greater surprises await us, and
I must admit that these examinations astonished me
greatly. In the drone antenna (Fig. 20) we have
thirteen joints in all, of which nine are barrel-shaped
and special, and these are covered completely by
smell hollows, before and behind, as at C, Fig. 22;
each hollow, beside, is somewhat less than those of
both queen and worker, being about yoVo m• in length,
an d y- Q - q 0 in. in diameter. An average of thirty rows
of these, seventy in a row, on the nine joints of the

ORGANS OF SPECIAL SENSE.

two antennae, give the astounding number of 37,800
distinct organs. When I couple this development
with the greater size of the eye of the drone, and
ask what is his function, why needs he such a
magnificent equipment? and remember that he has
not to scent the nectar from afar, nor spy out the
coy blossoms as they peep between the leaves, I
feel forced to the conclusion that the pursuit of the

Fig. 22.—Parts of Surface of Antenna (Magnified 360 times).

A, Portion of Front Surface of One of the Lower Members of the Flagellum (Worker
or Queen)—s', Smelling Organ ; Feeling Hair. B, Portion of Side and Back
Surface of One of the Lower Members of the Flagellum (Worker)—A, Ordinary
Hair; c'. Conoid Hair; A o (Auditory?) Hollows. C, Portion of One of the
Lower Members of Flagellum (Drone), Back or Front; lettering as before.
D, Portion of Lower Member of the Flagellum (Back, Worker or Queen).

queen renders them necessary, and that sight and
scent are the faculties by which this is accomplished.

The same wondrous little head that carries the
antennae, with their bewildering multiplicity of parts,
bears on its sides the extremely large compound,
or faceted eyes (Plate II., or A, B, Plate IV.). The
hand magnifier is sufficient to show something of
their structure, revealing that the beautiful satin-like
appearance they possess is due to their glistening
surface being divided into hexagonal convexities (H),
disposed precisely like the cells of honeycomb.
Each convexity, or facet, is little more than -j-g^in.
in diameter, and is, really, the outside of an indepen¬
dent instrument of vision. Between most of these

112

BEES AND BEE-KEEPING.

facets we find long, and generally perfectly straight,
hairs, which indicate, by their basal formation, that
they are sensory, as well as protective, in function.
The dark tone of the eye is due to the presence of
quantities of colouring matter (technically pigment)
within, and as this begins to form during the chrysa¬
lis condition, the growing eye then passes through all
shades between white and an intense purplish-brown,
approaching black. The external lenses, which are,
of course, devoid of colour, and very transparent,
are chitinous, being developed much as the external
skeleton, from an underlying layer, the latter disap¬
pearing when they are fully formed. Mutual pres¬
sure converts outlines which would, in its absence,
have been, in every case, circular, into a series of
hexagons—in proof of which, the lenses on the margin
of the eye (G) are bounded by a curve wherever
they are free; while those of the chrysalis are
circular, until they grow sufficiently large to bring
pressure upon each other.

If a specimen be properly prepared by hardening,
and then cut in cross section, the contiguous sets of
parts (ommatidia) are found almost in the form of a
fan (C). The superficial lenses, making up together the
cornea, are now seen to be bi-convex, while, beneath
each, is placed a second lens (the crystalline cone,
cc, C and D). (For the examination of the cornea and
crystalline cones, the directions at page 76 are suffi¬
cient ; but, for the finer parts, teasing with needles,
after soaking the optic tract for twenty-four hours
in a 5 per cent, solution of chloral hydrate, is neces¬
sary, unless staining and section-cutting, after har-

ORGANS OF SPECIAL SENSE.

1 T 3

dening, be adopted.) Beneath the crystalline cones
we have the great rods (rhabdia), consisting of several
straight chitinous threads, partly fused together
(gr, D), which pass inwards towards, and actually
perforate, the basilar membrane, which is represented
by a line running across the lower part of the fan¬
like form, in the section C. These rods are surrounded,
throughout their length, by eight retinulae, about
which are placed pigment cells, preventing the wan¬
dering of light from one optic element to another.
This will be best understood by the cross section
of the rods (F). The crystalline cones, for a similar
purpose, are protected by pigment cells, which have
also greater density at the upper, lower, and middle
portions of the rhabdia (marked ppp, C). Between
these microscopic telescopes, pointing in every direc¬
tion, run long and perfectly straight tracheal tubes,
which find their entrance by passing through per¬
forations in the basilar membrane. Immediately
behind the basilar membrane lies a complex nerve
structure, which Dr. Sydney J. Hickson,* in his
admirable paper, denominates the periopticon;
thence, running backwards, a bundle of optic nerve
fibrils (on, C), decussate (or cross), and then enter
a ganglionic swelling—the epiopticon, if we follow
the nomenclature of Dr. Hickson. Yet another
bundle of decussating fibrils brings us to the opticon
(g in our figure), beyond which lies the cerebrum,
described and illustrated at page 53. The structures
united by the decussating fibrils are complex, and

♦ “The Eye and Optic Tract of Insects.” The Quarterly Journal of
Microscopical Science, April, 1885.

BEES AND BEE-KEEPING.

difficult of examination. The periopticon is made up
of a number of cylindrical elements, seen just beneath
the basilar membrane (C), and consisting mainly of
divisions and sub-divisions of the decussating fibrils,
traversing a granular matrix, and for which structure
Dr. Hickson proposes the name of neurospongium.

All physiological students know perfectly that, in
our own eye, by example, vision, depends upon the
presence of nerve end cells, which lie behind the
expansion of the fibres of the optic nerve. The
retinulae, previously mentioned, have been very care¬
fully investigated by Grenadier,* and many other
naturalists, who have, with great unanimity, regarded
these structures as the nerve end cells of insects ; but
Mr. B. T. Lowne has recently published a treatise,t in
which he endeavours to show that the true nerve
end cells are situated behind the basilar membrane,
in the periopticon of Hickson. The controversy is
beyond our limits, and those desiring to follow it may
consult the works mentioned in the footnotes.

It is clear at once that the multitude of simple
eyes (directed to almost all points of the horizon),
which, by partial fusion, constitute the compound eye
of insects, permits a far wider range of vision than
would have been possible with a simple fixed eye;
but difficulties have been felt in explaining how these
parts produced a single true impression of surround¬
ing objects. Muller suggested that each ocellus saw
only the point just before it, and so a picture was

* “ Untersuchungen fiber das Sehorgan der Arthropoden.”

t “ Compound Vision, and the Morphology of the Eyes of Insects.”
Linnaean Society’s 7'ransactions, Second Series, vol. ii., Part n.

ORGANS OF SPECIAL SENSE.

; 5

made up in mosaic; but the remembrance of an obser¬
vation of Leuwenhoek showed the idea to be unten¬
able. The old Dutchman noticed that each corneule
(hexagonal lens of one ocellus) of a fly produced a
complete image of the flame of a candle, and not a
part of it. This introduces one of the most remark¬
able objects that the microscope can exhibit, and which
is quickly made from the eye of a bee (although
that of a beetle is better). Cut out the cornea,
wash it. clean inside with a camel-hair brush, place
it in water, under a cover glass, and use the flat side
of the mirror and a Jin. or Jin. objective. Focus
until all the hexagonal facets are visible. Now gently
draw back the objective, when, by daylight, a picture
of the window, with its bars, will be seen as formed
by each lens in the cornea. If the fingers be put into
the right position, their movements can be traced. I
have seen thus, simultaneously, in so many facets of
the bee’s eye, many hundreds of pictures of two
houses and surrounding trees, which stood 140yds.
from the microscope. At night, if a sheet of tissue,
with a watch face roughly drawn on it, be placed in
front of the microscope lamp, the time can be seen
through each corneule. The picture is alike in each ;
but then, it must not be forgotten that the cornea is
flattened by the cover, so that all the lenses are
made to look in one direction. The solution of the
question of multiple vision appears to lie in the fact
that each ocellus presents a slightly different picture
from its neighbour, since its axis is directed to a
different point, but that parts apparently overlapping
are identical, and so are interpreted into a picture by

L 2

BEES AND BEE-KEEPING.

the action of the ganglionic structures through which
the impressions pass. Those who have best studied
the most perfect eyes, know how true it is, that the
eye only looks, while that which lies behind it sees.

Besides the faceted eyes, bees carry three simple
eyes, called ocelli, or stemmata (I, Plate IV.), on the
upper part of the head, although they are not placed
quite similarly in the two sexes (see A and B). These
.eyes are very convex, and are adapted to short-
distance vision. Behind the simple lens (/, I), lie
structures much like those found in the compound
eyes. Indeed, these eyes, are posteriorly compound,
although anteriorly simple. It will be interesting to
note, by Fig. 12, page 53, and A, Plate IV., that the
cranium is so formed that the lateral ocelli should
have a range of vision sideways, while the middle
ocellus sees forwards, the hairs being so placed that
a clear outlook is preserved.

The possession of the colour sense by bees has
been well ascertained, and Sir John Lubbock’s* ex¬
periments have most satisfactorily shown, not only
a power to distinguish between, but a preference
for, particular tones. They have no doubt been, in
consequence of this faculty, active agents in develop¬
ing colour in blossoms, as we shall have occasion to
discuss in a future chapter ; while blossoms them¬
selves, by reaction, have played an important part
in augmenting their powers of discrimination.

The large space occupied in the head by the eye
structures has been strikingly shown by an interesting
case, recently brought to my notice, through th e
*“ Ants, Wasps, and Bees” (International Science Series).

ORGANS OF SPECIAL SENSE.

117

kindness of Mr. V. Novitzki, who sent me a number
of drones, the heads of two of which are accurately
drawn in Fig. 23. A is a true albino, the eyes, com¬
pound and simple, being alike absolutely devoid of
pigment. These drones evidently saw nothing clearly,
although they could distinguish light; for, if placed
in a box with a small opening, they found the latter
at once, and crawled out, but remained captives if

Fig. 23.—Heads of Abnormal Drones (Magnified Eight times).

A, Head of Albino— ce, White Compound Eye
Hairs; a, a, Antennse; m, Mandibula; mx,
Drone— h, Hairs; ]o, Outline, showing Size
a, a, Antennas ; m, Mandibula; mx, Maxillae.

0, White Ocelli ; th, Tactile
Maxillae. B, Head of Eyeless
ox Head of Normal Drone

the box was kept in the dark. In the same hive
with these drones appeared others with a still n30.ce
extraordinary defect—no eyes at all existing. The
owner represented to Mr. Novitzki that they were
headless, and such they might easily have been
supposed by a superficial observer. Smooth and
hard surfaces, bearing limp, irregular hairs, covered
the sides of the head, which did not rise high enough

BEES AND BEE-KEEPING.

to include the spot the stemmata would normally
have taken. In other respects, the head was perfect;
the antennae, jaws, upper lip, and tongue being well
developed. The dotted white line (B) shows the
space the eyes should have filled. Since albinism
is very uncommon in insects, I have sent specimens
of these bees to the Natural History Museum, South
Kensington, where they may at any time be seen.

It remains for us to compare the eyes of worker,
queen, and drone. Possibly, considerable variation exists;
so, for the purpose of comparison, I operated upon
bees from the same stock in each case. The worker
spends much of her time in the open air. Accurate
and powerful vision are essentials to the proper pro¬
secution of her labours, and here I found the com¬
pound eye possessing about 6300 facets. In the mother
of this worker I expected to find a less number, for
queens know little of daylight. After wedding, they
are out of doors but once, or at most twice, in a
year. This example verified my forecast, by showing
4920 facets on each side of the head. A son of this
mother, much a stay-at-home also, was next taken.
His facets were irregular in size, those at the lower
part of the eye being much less than those near the
top; but they reached the immense number of 13,090
on each side of the head. Why should the visual ap¬
paratus of the drone be so extraordinarily developed
beyond that of the worker, whose need of the eye
seems at first to be so much more pressing than his?
I have previously suggested the reason in considering
the antenna, but facts yet to come before us would
render further consideration of the argument premature.

ORGANS OF SPECIAL SENSE.

Some writers have described the eye of the

bee in a manner which seems to make all easy.
Their lenses appear to have been made in a lathe,
and run together by pigment which has been poured
in between them ; but this simplicity has the primal
defect of being inaccurate. The eye is gradually
evolved from elements incomparably more simple
than itself, and which existed in the blind larva. The
deep mystery of cell life has caused all to grow

into the form the mature eye possesses, but the
manner of its building is still dimly traceable—cells
still compose it, and the pigment is yet but a part
of the contents of some of these. Four cells, coming
together by the action of that life which is an ever¬
present miracle, by mutual action framed themselves
into the crystalline cone ; but the cone still shows
its origin, and is not like the homogeneous lens of

the optician; and so with every other part, for the

eye was and is vital. Every element made out only
leads back to some new and more recondite problem
yet to be faced. Can we, then, leave these sense
organs without being moved by their wonder ? Our
conception is unequal to the task they give us,
although our knowledge of them is, at the best,
only superficial. I feel unable to close this chapter
as I would. Swammerdam shall do it for me, for
he says : “ I cannot refrain from confessing, to the

glory of the immense, incomprehensible Architect, that
I have but imperfectly described and represented this
small organ ; for to represent it to the life in its
full perfection far exceeds the utmost efforts of
human knowledge.”

CHAPTER IX.

THORAX AND LEGS.

Centre of Movement—Simple Embedding—Freeing
the Thorax of Hairs — Legs: Insertion of — Joints:
Modifications of—Tarsal Hooks: their Uses — Pul-
villus: Uses of; How Cleaned; Automatic Action
of; How Folded — Front Legs: Brushes of —
Antenna Cleaner: Universally Possessed by Bees
—Second Leg Spine : its Use—Third Leg — Wax-
plate Pincer—Pollen Brush—Pollen Basket: Its
Structure; How Filled — Legs of Queens and
Drones—Droll Error—Comparison of Legs.

THE thorax, as the centre of locomotion, giving
attachment and movement to both legs and wings, is
necessarily nearly filled by large muscles; and these
are usually of a pink colour, as may be seen by
cutting a recently dead worker, or, better, a drone,
down the centre with a keen razor. For elementary
study, simple embedding, managed as follows, will be
very serviceable : Heat, only sufficiently to melt it,
some paraffin or bees’ wax, and place in this a few
bees. If the temperature be not too high, the longer
the “subjects” for dissection are kept soaking the

THORAX AND LEGS. 121

better. Roll up some wet writing-paper into the
shape of a tube, about ^in. in diameter, lift out the
bees with forceps, and drop them, one by one, into
it, so that they are arranged end to end. Fill up
the tube with the wax, and allow it to cool. The
paper being removed, the bees may be cut, in any
direction, into very thin slices, for examination.

The thoracic plates have their remarkable external
modelling completely concealed by the down which
thickly covers them above, and the long, webbed hairs
clothing them below. A small patch of these is seen
at I, Fig. 24, holding sundry pollen grains between
their meshes, the latter accomplishing their purpose, by
inevitably entrapping the granules furnished by the
anthers when visits are paid to blossoms. The
queen, as this would lead us to suppose, is relatively
bare beneath, but the drone is enveloped in a strong,
almost spiny, pubescence, giving him great clinging
power, of which the utility is apparent. A little
device will make the bees our assistants in studying
their thoracic and leg structure. Take a thin string,
about a foot long, and at each end fix a dead bee,
by tying round the neck. Drop the suspended
“ culprits ” between the frames of a stock, so that
the middle of the string rests like a saddle on the
top bar. In a couple of days, every hair will be
cleaned from the “ gibbets,” and their bodies polished
like those of beetles, so that the attachment of the
wings, the spiracles, the lines dividing pro-, meso-, and
meta-thorax, the actual form of the leg joints, and
the character of their articulations, with many other
interesting points, will be clearly visible. All adult

122

BEES AND BEE-KEEPING.

insects have three pairs of legs, which are inserted
into the three before-given divisions of the thorax.
Those of the bee, with their wondrous quickness

Fig. 24.—Drone and Queen Leg (Magnified Ten times), and Hairs Various.
A, Third Right Leg (Drone)— ti, Tibia; p, Planta, or Metatarsus; t. Tarsi.
B, Third Right Leg (Queen); lettering as before. C, Sensory Hairs from
Labial Palpus. D, Ditto from Maxilla. E, Split Hairs from End of Bouton.
F, Broken Compound Hair growing anew. G, Webbed Hair for Holding
Pollen Grains. H, Long Sensory Hair. I, Small Piece of Under Side of
Thorax, carrying Gathering Hairs and Pollen Granules.

and accuracy of movement, may be regarded from
two perfectly distinct points of view. First, as instru-

THORAX AND LEGS.

ments of locomotion, from which aspect the several
pairs, whether those of queen, worker, or drone, have
a common structure, and may be collectively, studied;
second, as supplying points, of attachment and move¬
ment to curious appliances, severally distinct in
character and purpose, and which, of course, require
individual treatment. Let us first examine them in
their locomotive capacity. The muscles moving them,
and which are energized as explained at page 51,
are partly within the thorax and partly within the
upper joints; while the lower ones carry only tendons,
moved as may be well understood by reference to
Fig. 10. Of course, in addition, the legs are provided
abundantly with both tracheae and blood. Each leg
consists of nine joints. Articulated into the thorax
we find the coxa, or hip (c, B, Plate V.), nearly
conical, and webbed beneath, and bearing the tro¬
chanter ( tr ), triangular, hairy, and firmly articulated
to the femur, or thigh (_/), which is the first elon¬
gated joint, and, like the previous ones, very densely
clothed by long webbed hairs. It is followed by the
tibia, or shank (ti), curiously modified in the different
pairs and sexes. A foot, or tarsus, of five joints, of
which the upper one (the metatarsus, p , Plate V.
or Fig. 24) is always much larger than the rest,
completes the limb.

The surprising power that bees possess, of suspend¬
ing themselves from the bodies of their companions,
and of sustaining a pull, without detachment, of many
dozens of times their own weight, which is rendered
apparent by the cluster formed in swarming, or in
the chains of workers festooning themselves from

124

BEES AND BEE-KEEPING.

the hive roof to its door, at the commencement of
comb building, is due to the strong claws, or anguiculi
(an, Fig. 25), which are found at the termination of
the tarsus. These claws, of great strength, bear a
secondary talon on the side, and carry long feeling
hairs ( fh ). They are capable of two movements.
They can be turned upwards, as at B, or point down¬
wards, as at A, Plate V. ; and, besides, they can be

Fig. 25.—Foot of Bee, with the Pulvillus in Use (Magnified Fifty times).
A, Under View of Foot— t, t, Tarsal Joints ; an, Anguiculi ; fh, Feeling Hairs ; pv,
Pulvillus; cr, Curved Rod. B, Side View of Foot; lettering as before. C,
Central Part of Sole— pd, Pad; cr, Curved Rod; fh, Feeling Hairs; pv,
Pulvillus Unopened.

made to approach each other, although not sufficiently
to meet. When turned upwards, the perfect support
they give to sisters desirous of forming another link
in the living chain is evident. By means of these
claws, bees walk on the edges of their comb cells, fix
themselves on the alighting-board, as they fan in the

THORAX AND LEGS.

135

summer sun, and cling on to the straw roof of their
skep, or the splintery roughness of their wooden hive.

But the tiny foot has yet another power ; for bees
can, like flies (although not with equal facility),
sustain themselves on the polished surfaces of leaves
and petals, and upon glass if needs be, although here
they get no fixing for their anguiculi. Placing a bee
in a bottle, and watching it through a hand magnifier
as it ascends the sides, we see between the claws a
whitish body (pv, A, Fig. 25), which seems to expand,
like the camel's foot, as the step is taken. I am not
aware that any observer has previously given the
pulvillus of the bee attention, yet it is so singular
and beautiful that to understand it is to be delighted.
All sorts of guesses (for guessing is so easy) have
been advanced to explain the fly's walk on the ceiling.
Some taught that its foot acted like a boy's sucker,
and that the fly was sustained by the pressure of the
air; but experiment quickly disposed of the error.
The fly can walk inverted on glass in a vacuum, but,
if it be moistened, the insect cannot walk on it at
all. So with bees : breathe on your glass super, or
manage so badly that it becomes moist inside, and
its surface altogether fails in affording foothold, for
the pulvilli give out a clammy secretion, which is
left in minute quantity behind, and I have found
high powers of the microscope to reveal its trace.
The moisture, of course, prevents this secretion from
taking effect. Dusting with flour, or very slightly
greasing, just as completely, makes perpendicular
smooth surfaces impossible of ascent. This will
explain why bees so object to plunging into pea

126

BEES AND BEE-KEEPING.

flour, when it is offered to them as artificial pollen
(see Artificial Pollen), and why, also, they so earnestly
clean their legs from all dust. The pulvilli are
cleared by rubbing the tarsi together, when the

pulvillus is drawn over their abundant hairs, which
are, in part, brushes provided for this very purpose.
Dredging flour over a bee will start at once this

movement, tiny pellets being dropped during the
operation, while the tongue is now and again out¬
stretched to supply saliva. Thus, the bee is able,

not only to clean itself, but to pack such a dry,

Fig. 26.—Bee’s Foot in Climbing, Showing Automatic Action op
Pulvillus (Magnified Thirty times).

A, Position of Foot in Climbing Slippery Surface, or Glass—pc, Pulvillus; fh,
Feeling Hairs; an, Anguiculus, or Claw; t, Tai-sal Joint. B, Position of
Foot in Climbing Rough Surface ; lettering as before. C, Section of Pulvillus
Just Touching Flat Surface— cr, Curved Rod. D, Same Applied to the Surface.

unadhesive substance as pea flour, in beautiful pellets,
on its hind legs.

We have seen that the pulvillus cannot be used
without loss of material. It is, besides, exceedingly
delicate, and easily injured by any roughness, so that
it is doubly desirable not to bring it into play where
the claws would take effect. I find all this is secured
by a most striking automatic arrangement. B, Fig. 26,
represents the pulvillus in its rest position, pointing
backwards, as it stands between the claws. If the
bee is ascending a rough surface, the points.of the

THORAX AND LEGS.

127

claws catch (as at C), and the pulvillus is altogether
saved from any contact; but if the surface be smooth,
so that the claws get no grip, they slide back, and
are drawn beneath the foot (as at A), which change
of position applies the pulvillus, so that it imme¬
diately clings. It is the character of the surface,
then, and not the will of the bee, that determines
whether claw or pulvillus shall be used in sustaining
it. But another contrivance, equally beautiful, remains
to be noticed. The pulvillus is carried folded in the
middle (as at C, Fig. 25), but opens out when applied
to a surface, for it has at its upper part an elastic
and curved, rod (cr, Figs. 25 and 26), which straightens
as the pulvillus is pressed down ; C and D, Fig. 26,
making this clear. The flattened-out pulvillus thus
holds strongly while pulled, by the weight of the bee,
along the surface, to which it adheres, but comes up
at once if lifted and rolled off from its opposite sides,
just as we should peel a wet postage stamp from an
envelope. The bee, then, is held securely till it
attempts to lift the leg, when it is freed at once;
and, by this exquisite yet simple plan, it can fix and
release each foot at least twenty times per second.

Space compels us to dismiss this part of an inviting
theme for the consideration of the legs as tool-bearers,
beginning with the front pair of those of the worker
(C, Plate V.). The pollen-gathering hairs, and the
soft skin, to admit of flexion between femur and
tibia, at once strike us ; while, upon the front of the
latter joint, we note a mass of close-set, soft hairs
(b), acting as a brush for sweeping the surfaces
which the coarser hairs have combed or scraped, and

128

BEES AND BEE-KEEPING.

for cleaning the semicircle of teeth presently to be
noticed. At the front of the metatarsus stands a set
of long, erect spines, which are always possessed by
those bees that have hairs between the facets of the
compound eyes. The spines ( eb ) are, really, eye-
brushes, or, perhaps, I should have called them
combs, since their office is to clear out from the
eye hairs all pollen grains or foreign bodies, which
would, of course, impede vision. But a most sur¬
passingly beautiful device of the first leg remains
to be noticed. It consists of two parts—a deep,
curved recess in the back of the metatarsus , and a
spine and sail, or velum ( v, C, Plate V.), attached
at the termination of the tibia. Professor Cook’s
reference to this marvellous mechanism is so inaccu¬
rate in every particular that he is best refuted by
quotation. He says: “On the anterior legs of the
workers, between the femur and tibia, is a curious
notch, covered by a spur. For several years, this has
caused speculation among my students, and has
attracted the attention of observing apiarists. Some
have supposed that it aided bees in reaching deeper
down into tubular flowers ; others, that it was used
in scraping off pollen; and still others, that it
enabled bees to hold on when clustering. The first
two suggestions may be correct, though other honey
and pollen-gathering bees do not possess it.” (The
italics are mine.) I must remark here, first, that
this appliance is not more possessed by workers
than by queens and drones; second, it is not between
femur and tibia, but where I place it; and, third,
it is possessed by every bee in this wide creation,

THORAX AND LEGS.

129

and also, in a modified form, by wasps and ants.
That the use of this appliance has been missed is
not astonishing, for the cause of failure is but too
evident. If bees be watched (and weary ones on a
window-bar are best for the purpose), the first leg
will now and again be raised in front of the head,
and then drawn outwards. The leg, by this move¬
ment, is put over the antenna, which slides up
past an especially-contrived slip-way, consisting of
the short, stiff hairs, near p in Plate V., until the
projection of the velum is reached, when the thread¬
like organ of hearing and feeling drops naturally into
the semicircular cavity. At once the tibia bends on
the metatarsus, and brings all into the position seen
at E; but the antenna is now compassed behind
and before, and the teeth of the semicircular comb
(standing up towards us in the Plate), as the leg
passes outwards, scrape off every particle of dust,
rendering all fit again for the delicate duties of
smelling and feeling. But the velum, too, aids in the
process. Its cross section ( v , H) shows a back pro¬
jection, which keeps the scraping edge in position
for its work, like the carpenter’s plane-body holding
the iron. The combs, made up of about eighty teeth,
of the form shown at F, are, of course, right- and
left-handed; the ends of their teeth, while engaged
in scraping, as at c, H, always going first. How
remarkable the device, and how exact the fitting of
parts! I have before stated that the queen’s antenna
is Y^-j-in. in diameter, and such is the measure of
the comb on her first leg; the drone’s, ^in., his
comb the same ; and so on among both hive and

i 3 o

BEES AND BEE-KEEPING.

wild bees wherever I have had the opportunity of
taking measurements. These antenna-cleaners are,
in all the genera and species, most charming objects
for low powers; and Mr. Enock has mounted many
at my suggestion, so as to display perfectly their
peculiar form. But we must pass on. The second
leg has no velum, but a conspicuous spur (.y) at the
termination of the tibia. This spur is the crowbar
by which the little forager levers out the pollen mass
(see page 18), which she carries home, stored in her
basket, seen opposite ti, at A. The second leg is
brought over the third, the spine enters at the top
end of the basket, and passes down behind the mass,
driving it forwards. This spine likewise aids in clean¬
ing the wings, and so is carried by both queen and
drone. The third leg is remarkably specialised,
and needs careful examination. The pollen-gathering
capacity of the hairs of the upper joints at B is
evident. The articulation between the triangular tibia
(ti) and oblong metatarsus (p) is quite at one angle
of the two joints, so that, as they move upon each
other, the parts opposite wp open and shut like a
pair of jaws, of which the upper is provided with
spiny teeth, shutting down over a flattened plate in
the lower. This nipper is exactly fitted to its
purpose, and is used for removing wax plates (soon
to engage our attention) from the abdomen of the
worker. Since neither queen nor drone produce wax,
the nipper is in their case absent (A and B, Fig. 24).

But the chief interest centres on the two joints last
mentioned, as a device for carrying pollen home to
the hive. The metatarsus is enlarged into a sub-quad-

THORAX AND LEGS.

rangular form, constituting a flattish plate, slightly
convex on both surfaces. The outer face (/, A, Plate
V.) is not remarkable, but the one next the body ( p ,
B) is furnished with stiff combs, the teeth of which
are horny, straight spines, set closely, and arranged
in transverse rows across the joint, a little project¬
ing above its plane, and the tips of one comb slightly
overlapping the basis of the next. Their colour is
reddish-brown ; and, entangled in the combs, we
almost invariably discover pollen granules, which have
been at first picked up by the thoracic hairs, but
combed out by the constant play of the legs over the
breast—in which work the second pair, bearing a
strong resemblance to the third, performs an im¬
portant part.

So soon as bees have loaded these combs, they do
not return to the hive, but transfer the pollen to the
hollow side of the tibia, seen at ti, A. This con¬
cavity, corbicula, or pollen-basket, is smooth and hair¬
less, except at the edges, whence spring long, slender,
curved spines, two sets following the line of the
bottom and sides of the basket, while a third bends
over its front. The concavity fits it to contain pollen,
while the marginal hairs greatly increase its possible
load, like the sloping stakes which the farmer places
round the sides of his waggon when he desires to
carry loose hay, the set bent over accomplishing
the purpose of the cords by which he saves his
property from being lost on the road. But a diffi¬
culty arises : How can the pollen be transferred from
the metatarsal comb to the basket above ? Easily ;
for it is the left metatarsus that charges the right

M 2

132

BEES AND BEE-KEEPING.

basket, and vice versa. The legs are crossed, and
the metatarsus naturally scrapes its comb face on
the upper edge of the opposite tibia, in the direc¬
tion from the base of the combs towards their tips.
These upper hairs standing over wp, B; or close to
ti, A (which are opposite sides of the same joint),
are nearly straight, and pass between the comb teeth.
The pollen, as removed, is caught by the bent-over
hairs, and secured. Each scrape adds to the mass,
until the face of the joint is more than covered, and
the hairs just embrace the pellet, as we see it in
cross section at G. The worker now hies homewards,
and the spine, as a crowbar, does its work.

Neither queens nor drones gather, and so their legs
are quite differently formed. The queen leg (B,
Fig. 24) shows, by its outline, that the worker is
a female; while the drone leg (A), rounded and
smaller, and not carrying even the rudiments of the
specialized hairs of the worker, is unlike either. An
explanation here becomes necessary, for it may be
remarked, that my drone and queen leg, according to
some authors, have changed places. It is so, but
for the following reason : An old French entomologist
published some capital drawings of bees' legs, but
his numbering read backwards, since the revers¬
ing action of the printing press had been forgotten.
He was copied by Blanchard, who, failing to note
his authority's mistake, called the drone leg the queen
leg, while the latter went to the credit of the drone.
Dr. Duncan translated Blanchard* and, quite inno¬
cently, and very pardonably, repeated Blanchard’s

* “The Transformations of Insects.” Cassell, Petter, and Galpin.

THORAX AND LEGS.

; 33

blunder; while Cook, who has taken many of the illus¬
trations in Part I. of his Manual from Duncan, con¬
tinues to ' the present hour to publish the error.
Surely, after eleven editions, the time has come for
breaking the spell, and giving the queen her own
legs back again. But, seriously, it is a pity when
authors become so fashionable as to slavishly follow
the antique. Doubtless, they are saved much time
and trouble; but their readers are wronged if they
are made to devote their time to obsolete fiction,
and unchecked mistakes, when they are led to
believe they are studying modern research.

I shall not again refer to Professor Cook’s book,
nor should I have done so at all, had not the
interest of scientific apiculture demanded it.

In comparing the legs of queens, workers, and
drones, it is worthy of remark that the queen has by
far the largest set, as she is a great walker, constantly
perambulating the combs. The drone depends little
upon his legs, and so he, notwithstanding his greater
weight, carries smaller ones even than those of the
worker. They are also but little specialised, their
principal peculiarity lying in the hairs of the smaller
tarsal joints (/, A, Fig. 24), which, in his case, are
heavy, and, instead of being simple, are strongly
webbed, so as to assist him, as indicated at page
121. The curious adaptations already observed, where
none were formerly suspected, makes it certain that
future investigations must greatly increase our ad¬
miration of such an inconsiderable matter as the leg
of a bee.

CHAPTER X.

WINGS AND FLIGHT. BUZZING AND HUMMING.

Development of Wings: Nervures, Cells, and Hairs

— Wings in Diptera : Reasons for Four Wings in
Bees and other Hymenoptera — Hooklets—Posterior
Wing not Flat — Comparison of the Wings of
the Sexes—Drone can Fly Backwards—Bee Line —
Flapping Movement Converted into Flight: Its
Velocity — Experiment■—Forward Flight—Backward
Flight: its Necessity ; how Performed—Ascending
and Descending Flight — Steering—Wing Rate,
Graphic and Musical Determination of—Buzzing
and Humming — Obturator Apparatus — Tracheal
Distension—Specific Gravity—Sonorous Membrane

— Voice.

The four membranous wings of hymenopterous insects,
articulated in pairs into the meso- and meta-thorax,
are formed in the chrysalis from vesicles, or
flattened -pouches, extravasated or pushed out from
the epidermal layer (see Fig. 4), and which are
brought into form by a series of interior tubes of
chitine, called, in the mature organs, nervures, and
seen, in Fig. 27, to divide both anterior and pos-

WINGS AND FLIGHT.

*35

terior wings into cells. The entomological names of
those of the anterior wing are given with the illus¬
tration, as they are frequently used as a basis of
classification.

When, by re-absorption of the contained nutrient
fluid, the two facing membranes of each flat pouch
are intimately joined, they become the transparent
extension of the wing, stretched upon the nervures,
which form its stiffening framework. The hollow
nervures are never wholly deprived of blood, while
through them run large tracheae, which, at the exit of
the bee from the cell, aid it in giving that full expansion
to its new organs which their office demands. As the
eye has left upon it the marks of its method of develop¬
ment, so the wing gives traces of its origin. The
microscope shows that it is dotted over on both sides
by small, stiff hairs with an expanded base, while very
careful examination reveals that the whole surface is
divided, by faint, angular lines, into small areas, which
indicate the boundaries of the primary cells, upon the
middle of each of which stands a single hair.

Every wing—be it of bat, bird, or insect—that is
capable of acting effectively as an instrument of flight,
must, in area, bear some definite proportion to the
weight of its possessor. The common bluebottle, a
dipterous insect, somewhat less than the honey bee,
has its single pair of wings of such a width and so
placed that their points are -|in. apart when at rest.
Had the bee been similarly formed, its wings would
have barred its entry to its own cell, which is only -jfin.
in diameter ; so that cleaning, filling, and emptying of
comb, feeding of brood, and many other essentials,

BEES AND BEE-KEEPING.

would have been impracticable. This difficulty, how¬
ever, is exquisitely met by the necessary wing-surface
being made up by two pairs, an anterior and a posterior,
which lie one over the other in repose, so that they
occupy but little space, their two points in position

A and B, Anterior and Posterior Right Wings of Worker (under side), Magnified
Eight times—1, Costal Cell; 2, Externo-median Cell; 3, Interno-median Cell;
4, Anal Cell; 5, Marginal Cell; 6, 7, 8 and 9, 1st, 2nd, and 3rd and 4th Sub¬
marginal Cells ; 10, 11, and 12, 1st, 2nd, and 3rd Discoidal Cells ; 13 and 14,
1st and 2nd Apical Cells ; c, d, Plait; e, /, Hooklets. C, Plait and Hooklets,
Magnified Twenty-five times— c', d', Plait; e 1 , Hooklets. D, Cross Section
through line a, b, of p. Plait, and A, Hooklet, locked together.

only covering a width of fully ^-in. Other hymeno-
pterous insects have, in this respect, a like structure,
and for identical reasons : the ant travelling through
narrow galleries, the wild bee in its burrow, and the
wasp in its cell, being able to so place their wings

WINGS AND FLIGHT.

137

that they offer no impediment to their home move¬
ments, while the neatness of their packing is in itself
a security against damage. The queen of the bee¬
hive, indeed, proverbially carries her wings very
closely set over the back (see Fig. 5), for the
greater length of her life demands the greater care ;
and so the gauzy membranes, in her case, are capable
of sustaining the wear of three or four years, yet re¬
maining good enough for duty.

Presently we shall discover that the rate of vibration
given to the wings during flight is prodigious, and then
the division, so valuable during repose, becomes an
impediment, for the air cannot be so efficiently beaten
by two narrow wings as by one of their united width.
And here, again, a device, charming in its mechanical
simplicity and perfection, presents itself. The inner
margin ( c , d , A, Fig. 27) of the anterior wing is folded
under, in a plait, while a series of minute blunt hooks
( e , f, B) are turned up upon the outer margin of the
under, or posterior one. As the anterior (upper)
wing moves outwards into position for flight, its
down-turned plait passes over the upper surface of
the lower wing, and is caught by the upturned
hooks, as C and D will make clear; and now the
two wings, wedded into one, strike the air: but, at
the moment the flying insect settles, these, by falling
back into position, become immediately free, since the
plait simply slips from the hooks, and the wings take
up their superposed position.

The hooklets decrease in size in beautiful grada¬
tion towards the wing point—the largest are about
-jig-in., the smallest, in length—but they are not

BEES AND BEE-KEEPING.

138

always the same in number on the two sides of the body.
The posterior wing is, most suggestively, not absolutely
flat, but convex above, in the direction of its length,
so that its hooklets are held up towards the plait
on the anterior wings, the hairs just behind which
turn in a direction different from the rest, so that
the movements of the hooklets shall meet with no
impediment. How well Nature rewards looking into
even the smallest matters!

A comparison of the sexes is again instructive.
The queen is commonly said to have smaller wings
than the worker. Yet this is only true relatively,
and clearly for the reason that she has much less
frequent use for these parts than her ever busy
children ; but the drone, lighter than the queen, is
endued with that soaring power and rapidity which
his function renders necessary, he possessing organs
of flight far larger than hers, and which extend
beyond the extremity of the abdomen (see Fig. 5).
The measurements are given in one-hundredths of
an inch.

Length of Length of Ratios of

Anterior Wing. Posterior Wing. United Area.

Worker.38 .... 28.5

Queen.41 .... 29.6

Drone.49 .... 35.9

The remarkable strength and width of the inferior
wing of the drone gives an intimation, which
observers should keep in sight; for we shall see
presently that this enables him to fly backwards with
great energy, should such a necessity arise. The
relative perfection of the organs is well indicated
by the hooklets; and here, again, we find the drone

WINGS AND FLIGHT.

139

in the van, and the queen in the rear. The queen’s
hooklets vary considerably in number, ranging from
thirteen to twenty-one; the worker’s, nineteen to
twenty-three; the drone’s, twenty-one to twenty-six.

Bees are accomplished fliers, but they never traverse
the air with the same directness as many birds, so
that the expression "bee line,” used by bee-hunters,
needs to be accepted in a modified sense. It is their
habit to skim along, in extended sweeps, alternately
curving to right and left. The rapidity of their aerial
voyages is difficult to calculate. Stories have been
detailed of their darting in and out of the windows
of a train, in rapid movement, but these furnish no
evidence of their velocity when unaided, since the
train carries the air lying in its neighbourhood along
with it, as leaves and paper scraps frequently make
clear. My own observations lead me to suppose
that the pace ranges between two and sixteen or
eighteen miles per hour, depending upon the load
and the nature of the errand—a bee, bearing the
body of a deceased sister from the hive, taking
the funereal pace, while those issuing forth on
business bent go express.

We must now turn our attention to the means
by which the mere flapping movement of the wings
is made to translate the creature through the air,
forwards or backwards, at any velocity less than its
maximum, and in any direction it may desire.

Fig. 27 shows a strong chitinous rod, called the
costal nervure, running along the anterior margin of A;
and it is this nervure, carried up and down by the
reciprocal contractions of the depressor and levator

140

BEES AND BEE-KEEPING.

alarum muscles, which moves the membranous exten¬
sion lying behind it constituting the wing.

A simple experiment, which I would recommend
all my readers to try, and which I have often used
as an illustration, will make clear at once how this
arrangement wafts the insect forwards ; but we
must be careful to remember, in interpreting it, that
the wings in flying are not . carried over the back,
but are brought round, with their length approxi¬
mately at right angles to that of the body, so
that the costal nervure goes first, and is followed
by the membrane. Gum or glue the edge of a

pw, Paper Wing ; s, Stick; DS, Down Stroke ; US, Up Stroke ; c, e, e, Air Currents.

piece of writing-paper, 3m. or 4m. wide, along
the stick of a penholder, or some such form, so
that the paper represents the wing-membrane, and
the stick the costal nervure. Now place two
lighted candles as in Fig. 28, and wave the paper
up and down between them, so holding the stick
that while it is at rest the paper is horizontal.
Both flames will immediately indicate a current from

WINGS AND FLIGHT.

141

the stick towards the paper slip. When the down-
stroke (DS) is made, the resistance of the air throws
the paper relatively up, and the air is reflected

from its surface, as indicated by the arrows. Simi¬
larly, when the upstroke (US) follows, the paper is,
by resistance, thrown into such a position that the
air is reflected in the same direction as before, so
that both ascending and descending strokes give
an identical current. Simple mechanics shows that
the current from right to left in the Figure, by re¬
action, tends to move the paper and stick from left
to right. Applying this now to the bee, whose

pliant wing-membrane yields to pressure like the
paper, we learn that both up and down strokes pro¬
duce a current towards the costal nervure, and from
the posterior edge; or, in other words, that the bee’s
wing itself is moved in space, the costal nervure

going first— i.e., the bee flies forward. It is un¬

doubtedly interesting to thus note how both up and
down movements aid in progression in one line. Yet
this fact but opens up another inquiry, for, if the
bee were only able to fly forwards, her plight in its
measure would resemble that of a steamship which
could not reverse her engines ; they might be stopped,
but she would remain under weigh, to possibly com¬
pass her own destruction ere her initial velocity had
become expended. But a little attention in an apiary
will make evident that bees are competent to wing
their course backwards. As young ones come out
for their first airing in the warm mid-day sunshine of
spring, they fly constantly looking to the hive door,
advancing and receding in curves, so that the head

H 2 BEES AND BEE-KEEPING.

frequently follows the body. If a bee be watched,
too, honey-gathering— eg ., on an apple-tree—she flies
rapidly from flower to flower ; but, at the exact
moment, her hasty advance is suddenly and mysteri¬
ously checked, so that she plies her quest by a touch
of such measured delicacy, that no filament, however
tender, is broken, and no petal unduly pressed. But
by what means is this sudden stopping, or this back¬
ing, secured? And here we get a deeper insight into
the meaning of the small wings than that previously
gained ; for, although it is clear that they consider¬
ably aid in sustaining the bee, from the fact that
she can fly down, but not up, after their removal,

Fig. 29.— Section op Wings (Magnified Twelve times), to Explain how Flight is
Directed.

aw, Anterior Wing; pw, Posterior Wing; en, Costal Nervure; p, Plait; h,
Hooklets; c, Air Currents; H, Position of Head; A, Position of Abdomen ;
DSF, Down Stroke, Flying Forward; DSB, Down Stroke, Flying Backwards.

yet they subserve other purposes, by adaptations which
cannot fail to strike us as unspeakably beautiful.

In Fig. 2 8 we observe that the up and down move¬
ments of the stick are wider in range than that of
the paper, and that its motion decreases as we pass
from right to left; similarly, in Fig. 29, where the
cross section of the two locked wings is given, at
DSF (representing the position at the down stroke
when flying forwards) we must note that the large
wing has a more extended beat than the smaller, since
the latter is the equivalent of that part of the paper

WINGS AND FLIGHT.

[ 43

lying between a and b, Fig. 28. But should the
bee desire to reverse her movement, decreasing the
energy of the larger wing, and increasing that of
the smaller, instantly accomplishes her purpose, with¬
out any stoppage of flapping, because the then
stronger beat of the small wing, and the restricted
beat of the larger, immediately reverses the set of
their united plane. By examining DSB, Fig. 29,
we shall see the truth of this. Here the wider move¬
ment of put) (the posterior wing) makes it the leader,
producing the alteration that would have arisen from
transferring the stick to the opposite edge of the
paper (Fig. 28), and the air is, in consequence,
beaten in the opposite direction ( c '), so that the
bee is carried backwards, abdomen first. The up
stroke, as before, producing the same current as
the down, another Figure is not needed to repre¬
sent it. The case is that of a screw steamer
which, without stopping her engines, reverses her
course by changing the direction of the pitch of the
arms of her propeller. This, however, is my theory,
as distinct from that of Gelieu, which is immensely
more complex, and would require a nervous control,
which seems to me utterly incredible.

The question of ascending or descending now
suggests itself. This has been fully investigated by
Marey,* by means of adjustable models, of which
our space will not permit a description. Observations
on bees themselves have led me to the following
conclusions : The wings, during flight, are the points of

* “ A Treatise on Terrestrial and Aerial Locomotion ” (International
Science Series).

144

BEES AND BEE-KEEPING.

support beneath which the centre of gravity, if free
to move, always arranges itself. If a ball be held up
by a string, the centre of gravity (identical with the
centre of the ball, if the latter be of uniform density)
comes to rest under the point of support, towards
which position it immediately falls after every dis¬
turbance. If, in flight, a bee desires to rise, she
straightens out her abdomen, thus carrying her
centre of gravity (or weight) backwards, and, as a
consequence, the abdomen, with regard to the rest,
sinks, and the head points upwards, the body
revolving around the wings, so that the before-men¬
tioned centre of gravity occupies a position beneath
them. This alteration in the direction of the body
makes the flight one of ascent. But, on the con¬
trary, curling the abdomen beneath, by the action
of muscles lying in the thorax under the meso-
phragma, brings the centre of gravity forwards, and
allows the head to relatively sink, and a descent
in flight is the result. It is possible that the ab¬
domen, acting after the manner of a rudder, also
occasions, similarly, all lateral changes of direction.
It certainly partially effects these movements, but
whether assisted by inequality in energy of the right
and left wings, steering as the sculler does, is not
yet determined.

The marvellous velocity with which the wings of
most insects vibrate has excited considerable atten¬
tion, and has been tested by most ingenious experi¬
ments.

Let us first refer to what is known as the “ Graphic
method.’' A metal drum, revolved by clockwork, is

WINGS AND FLIGHT.

: 45

surrounded by smooth paper, which has been coated,
by exposure to a smoky flame, with a thin and easily-
removed sooty deposit. If a living insect be so held
that the wing in vibrating just touches the paper
while the drum is rotating, a series of scratch-like
marks, equi-distantly placed ( c , d, Fig. 30), will
indicate, by the spaces between them, the amount of
movement made by the drum during the time occu¬
pied by each vibration of the wing. This time is
accurately determined as follows: A tuning fork,
whose note (and, consequently, exact number of vibra¬
tions per second) is known, has one of its prongs

Fig. 30.— Graphic Representation op Rate of Vibration op Bee’s Wing.
a, b. Line Made by Tuning Fork; c, d, Marks Made by Vibrating Wing.

provided with a small pointed style. The latter, at the
moment the insect is being operated upon, is brought
into contact with the revolving drum (the fork, of
course, sounding), and is so held that the style moves
up and down upon the sooty paper. A waved line
(like a, b, Fig. 30) is produced, the length of each
wave marking the space traversed by the drum while
the fork makes one vibration. Should the fork give
256 vibrations in a second (sounding the middle C),
256 waves will occupy the space moved through by the
drum in the same time. If opposite to these should
stand 190 dots made by the bee’s wing (c, d, Fig. 30),
we get 190 vibrations per second as its rate—the

146

BEES AND BEE-KEEPING.

result at which Marey arrived. Tremendous as this
speed appears, involving a sequence of muscular
contractions of almost inconceivable rapidity, it is
probable that it is considerably below the truth, both
because of the weakening effect of the experiment
and the friction of the paper; Marey finding that,
as he lessened the contact of the wing on the drum,
the velocity very considerably increased.

These objections do not attach to determinations
based upon the note the flapping wings produce.
From what has been said of the tuning fork, it will
be remembered that pitch depends upon the number
of vibrations in a given time, and as the note
formed by the wing of the bee in vigorous flight,
according to Landois,* ranges between the A and
C of the first and second ledger of the treble clef,
its velocity, if this musical determination be accu¬
rate, can be no less than about 440 vibrations per
second, instead of 190, as reached by the Graphic
method ; but Landois himself observes that fatigue
has a marked effect, quickly bringing the rapidity
down to three-fourths of its normal amount.

In this connection it is worthy of remark, that
bees in the full vigour of youth and health are not
always in a condition in which flight is possible.
They may, now and again, be noticed to content
themselves with running , although frightened, even
touches with the finger at first inducing no more
than flying jumps of 3m. or 4m. Their temporary
inability is due to the small amount of air the

* “Die Ton-und Stummaparate der Insecten. Zeitschrift fur Wissen-
schaftliche Zoologie, 1867,” page 105.

WINGS AND FLIGHT.

tracheae contain at the time. They are at rest, the
blood is moving slowly, the body is specifically heavy,
and the muscles are not braced up ; ^mt after the
wings have been lifted, and a few energetic move¬
ments of the abdomen made, the vesicles and tracheae,
which just before were flat as ribbons, get filled,
and the bee sails away. In many practical opera¬
tions, bees may be shaken down from their combs in

Fig. 31.— Longitudinal Section through Thorax of Drone
(Magnified Seven times).

LA, Levator Alse (Wing-raising) Muscle, showing Fasciculi, or Fibre Bundles;
DA, Depressor Abe (Wing-lowering) Muscle ; A, Antagonist of Depressor ■
pwm. Posterior Wing Muscles ; mp, Mesophragma ; as, Air Sacs ; No. 3, Gland
No. 3; c, Cervical or Neck ; A,.Part of Head.

a mass, scooped up in spoons or shovels, and weighed
and measured in open vessels, pretty much like seeds ;
the facts just recounted going far to explain the reason.
The utility, beyond the purposes of flight, of filling up
with air, and the method of its accomplishment, are
both interesting and curious. Fig. 31 gives a section
through the thorax of the drone, showing the muscles
of flight, surrounded on all sides by air sacs (as), from
which pass very numerous tracheae (page 42), supply¬
ing the abundant oxygen these most active muscles re-

BEES AND BEE-KEEPING.

quire. As DA (the wing-depressors) contract, they pull
the mesophragma ( mp , and page 88) forwards and up¬
wards, and away from the metathorax. The separation
of the two walls of the air sac lying behind the
mesophragma draws in a supply of air, which, at the
relaxation of the depressors, is distributed to the
tracheae, as the antagonist muscles (A) replace the
mesophragma, and rapidly drive all the air from the
air sacs. Other movements aid in the work, so that
the initial efforts of flying, as a natural result, distend
the body, and bring about all the conditions the absence
of which we just now noticed as making soaring im¬
possible. We shall presently see that the bee has
perfect control over the spiracles, closing them at
pleasure. When on the wing, then, with the air sacs
fully filled, if the spiracles be shut, the power is
gained for discharging the contents of the bowels by
simple pressure, the latter being applied by con¬
traction of those muscles which govern the abdominal
rings. That bees labour without weariness in banish¬
ing every vestige of impurity from their hives, which,
under natural and healthy conditions, they never soil,
has frequently been remarked. But these most cleanly
creatures are, in this latter respect, structurally com¬
pelled so to be, from the above-given curious arrange¬
ment. The queen is an exception, so far as her
capability of removing the intestinal residua is con¬
cerned, as her ovaries occupy the space taken by a
pair of large air sacs in the worker and drone; so
that she on foot, and for an obvious reason, possesses
the power (pages 71 and 84) the others only acquire
when , on the wing.

BUZZING AND HUMMING.

Every practised apiculturist knows that both workers
and drones emit a tone during flight, which is subject
to considerable variations, and that these often furnish
some indication of the particular “ frame of mind ”
of the insect at the moment. The reasons for some
of these differences have already been hinted, but it
would be extremely erroneous to conclude that the
wings alone, or even mainly, give out a note, in proof
of which an easy experiment may be cited. If one
of the larger Humbles— e. g., Bombus terrestris or
hortorum —be shut in a box, after removal of the
wings, or after they have been so gummed as to be
incapable of movement, a humming note will still be
produced, which, under the excitement of fear or anger,
may be even violent; anatomy showing that this sound,
which accompanies the true tone of flight, results from
a membranous vibration in the spiracles, the latter
being, amongst honey bees, especially developed in the
drone, whose sonorous qualities were referred to by
Shakespeare. Landois, to whom reference has pre¬
viously been made, recognised three tones in the flight
sound : the first, produced by the wing beats; the
second, sharper in character, by the vibrations of the
abdominal rings ; the third, the most acute and intense,
from the action of the true vocal apparatus, placed in
the stigmatic orifices. He found that stopping these
orifices with wax brought the humming to an end at
once. The wings undoubtedly do the buzzing, but the
humming is as clearly the outcome of an apparatus
formed as follows : The spiracles (page 33) have each
lying behind them, in a vestibule (or sounding-box)
made by an enlargement of the commencement of

BEES AND BEE-KEEPING.

* 5 °

the tracheal tube, a chitinous ‘‘stirrup,” or crescent¬
shaped piece, the object of which is to give the
insect the opportunity of voluntarily closing the air
openings, and this for a before-mentioned reason. A
double lever, formed of two irregular and unequal
cones, and actuated by an obturator, or closing muscle,
and tendon, is so contrived that the contraction of
the muscle causes the plugging of the trachea open¬
ing out of the back of the vestibule. The sound is
actually emitted by curtains, somewhat plaited and
fringed, formed from folds of the membrane lying
behind the edges of the spiracle, and in front of the
stirrup and lever. Muscular contractions within the
thorax, occasioning the wing vibrations, rapidly puff air
in and out, and so start the curtains in producing
that hum, which varies according to their tension,
and which may not inaptly be called the bee’s voice,
since it results from the movements of an apparatus
not unlike that of voice in ourselves and the higher
animals.

How many wonders are involved, then, in simple
flight ! The floating of the little insect, as it plays in
the sunbeam, or the rapid transport of it at plenty’s
distant call, enabling it to round a thousand corners,
and drop with the greatest accuracy into the mouths
of countless flowers, with the wafting of it back
again to its desired haven, singing, as it goes, from
many mouths, is not accomplished without the framing
of a mechanism which is all worthy of our admira¬
tion, and which has actually excited the envy, whilst
it has mocked, and is mocking, the inventiveness
and resources, of mankind.

CHAPTER XI.

SECRETION OF WAX, AND BEE ARCHITECTURE.

Reaumur’s Hypothesis—The Discovery — Dr. J. Hunter
—Frangois Huber and Burnens — Wax Pockets—
Wax-yielding Membrane — Queen and Drone —
Microscopic Examination — The True Gland : Its
Structure—Wax at First a Fluid—Wax Scales —
Wax of Comb — Wax Produced from Saccharine
Substances, not Pollen — Huber’s Experiments —
Comb Building Exhausting—The Behaviour of a
Swarm—Conditions Favourable to Producing Fat
and Wax—Structure of Comb — Model Making —
Cramer’s Demonstration —Angles — Economy in
Space—Equality of Solid Angles — Maraldi’s Calcu¬
lation—Strange Myth—Bees Capable of Modifying
Comb—Suitability of Hexagons — Soap-bubble Ex¬
periment—Costliness of Wax—Queen Cell—Covers
of Honey not Air-tight; Why Irregular—Sizes
of Cells—Drone Cell Peculiarities—Breeding Cells
always Approaching the Ideal Form.

The opinion formed by Swammerdam and Maraldi,
and accepted by Reaumur, that pollen, which he called
crude wax (page 16), was submitted to a peculiar
elaboration in the stomach of the bee, whence it was

152

BEES AND BEE-KEEPING.

returned to the mouth as true (“ veritable”) wax, was
completely overthrown by a French peasant,* whose
name, unfortunately, has not survived, he discovering,
in August, 1768, that the substance used in the con¬
struction of comb emanated, from between the rings
of the abdomen. This humble inquirer, a member of
a society of bee-keepers founded in Lusace even at
this early date, appears, after having pulled some bees
from comb they were then building, to have removed
their wax scales by the aid of a needle ; but his
pregnant observation slumbered for twenty-four years,
when Dr. John Huntert partially investigated the
subject, and drew attention to the existence of wax
glands. In the following summer, the blind Francois
Huber, justly admired for his researches, and deserving
the honour of all good men for his noble acknowledg¬
ment of the immense help he received from his ser¬
vant Burnens, repeated the discovery of the peasant,
and entered upon a series of experiments and observa¬
tions which will keep his name green so long as
apiculture is practised.

We already know that the abdomen of the worker
is arranged in six dorsal and six ventral inelastic
plates, which may move upon each other, because
they are united by delicate membranes, giving to
the whole the arrangement of the tucks of a child’s
frock. The exposed part of each ventral plate is
tough, and covered by webbed hairs ( wh , Fig. 32),
which much decrease in size towards the anal ex-

*Langstroth makes a prior claim for Hornbostel, in 1745, but he
gives no details of what was seen.

f “ Philosophical Transactions,” 1792.

WAX, AND BEE ARCHITECTURE. 153

tremity ; but if the abdomen be elongated by gentle
traction, we begin immediately to catch sight of ex¬
tremely smooth and delicate expansions (W, W), upon
which, very generally, in the warm season, wax plates
of greater or less size and thickness may be dis¬
covered. These pale yellow tender discs, which have
sometimes been called, quite incorrectly, the wax
glands, are eight in number, being found on the

Fig. 32.— Abdominal Plate (Worker), Under Side, Third Segment
(Magnified Twenty times).

W, Wax-yielding Surface, covering True Gland ; s, Septem, or Carina; wh, Webbed

four ventral plates intervening between the first and
last. They are surrounded and held in position by
a frame-like thickening of the plate itself (Fig. 32),
while between them runs a septem, or carina (3-).
The contour of the membranes determines the form
of the wax scales, which are moulded upon their
surfaces as the secretion passes, by osmose from the
true glands beneath. The hinder part of each

r 54

BEES AND BEE-KEEPING.

ventral plate covers the membrane of the ring next
it, forming with it a little pouch (wax pocket), open¬
ing backwards, from which the wax scales often
protrude a, considerable distance. The queen and
drone, on the under side of the abdomen, are in
this respect quite differently formed from the worker,
wax glands being entirely absent in their case, since
they take no part in comb building. The plates of
the queen (B, Fig. 33) are wide, to give her greater

Fig. 33. -Abdominal Plates, Under Side, Third Segment (Magnified
Twenty times).

A, Plate from Drone—a, strap ; 6, Webbed Hairs ; sh, Short Hairs. B, Plate from
Queen ; o', Strap ; c, Down ; sh, Short Hairs.

length of body, while the webbed hairs are wanting,
since these would interfere with ovipositing, and no
carina exists. The corresponding plate (A) in the
drone, though strongly framed, is narrow, because his
abdomen carries seven rings instead of six. Loose but
stout webbed hairs are provided, for reasons previously
noticed, and the process (a) giving attachment to
muscles aiding in abdominal contraction is much
stronger than that (a') possessed by the queen.

WAX, AND BEE ARCHITECTURE.

Examining, by a medium power of the microscope,
the wax-yielding surfaces, as removed from a bee’s
body, an appearance is presented not unlike that of
B (Plate I.) ; but this is due to an underlying single
layer of cells, which, by mutual pressure, are driven
into irregular hexagons. After carefully removing the
cells just mentioned, I find no evidence of structure
in the discs, although, by their character of fracture,
they are shown to be double, or to consist of dense
faces, with softer material between. The cells, of which
there are about 140,000 in the eight glands, when in
situ are very closely applied to the external discs

Fig. 34.—Portion of Wax Gland, seen from the Side Bathed bt Blood
(Magnified 800 times).

tr, main trachea ; n, nucleus ; o, o, o, oil-like globules.

(W, Fig. 32), at whose edges they most abruptly ter¬
minate. They collectively form the true glands, are
each about —g-t^in. in diameter, and contain a large
nucleus and many small granules, the latter occasionally
in movement; besides these, some of the cells seem
almost filled up with oily-looking globules [0,0,0, Fig. 34);
and it is also remarkable, that the part of the surface
of each cell which lies next the membrane is raised
into numerous minute prominences, needing for their
detection careful illumination and the highest order

i 5 6

BEES AND BEE-KEEPING.

of objective. The greatest peculiarity of this cell-
layer consists in the arrangement and abundance
of its small tracheae (Fig. 34), which do not pass
over the upper or lower surfaces of the cells, but
travel between their contiguous walls, in such vast
numbers, and with such repeated loopings, that con¬
stantly as many as five or six interpose in a space
which cannot be greater than the aoV oth of an
inch. The larger tracheae (tr), supplied from the ab¬
dominal air sacs, divide into finer ones, which imme¬
diately plunge between the cells, and there take a
course which, in the aggregate, amounts to about 60ft.
in length. This great need of oxygen for wax secretion
is highly suggestive, and will make clear a difficult point
when treating hereafter of the chemistry of the hive.

Wax, like every secretion, vegetable or animal, is
at first liquid. It is derived from the blood by cell
action, and then, transuding the structureless membrane,
assumes the solid form of the scale, which, if lifted
when the gland is active, will always show that it is
fluid beneath. While examining this question, I was
struck by finding that the webbed hairs ( wh , Fig. 32)
had their webbings in part or wholly covered by a
perfectly fitting casing of pure wax, which could only
have arisen by a transference of the secretion, while
still fluid, to their surfaces.

Turning our attention now to the wax scales, we find
them to differ from the wax of comb. They are much
more brittle and transparent, being not unlike flakes
of talc. Turpentine dissolves them immediately with¬
out residue, whilst fragments of comb disappear but
slowly in the same medium, which they make cloudy.

WAX, AND BEE ARCHITECTURE.

*57

Ether melts wax with difficulty, the scales for a long
period remaining in it intact ; whilst comb-wax
breaks up into minute fragments. When the bee
is engaged in building comb, the wax scales standing
out beneath the pockets, as we see them in Fig. 35,
are removed, as required, by the pincer of the third
leg (page 130), which is applied immediately against
the body, with the planta (/>, B, Plate V.) turned
from the tibia, so as to widely separate the jaws
of the pincers, whose bristle teeth are now passed

Fig. 35. —Under Side or Worker, carrying Wax Scales
(Magnified Three times).

adroitly beneath the wax scale. The two joints
being brought into line, the teeth pierce the scale,
which the leg in turn draws from the secreting
membrane, to be transferred to the front legs, and
thence to the mouth, where it is held perpendicu¬
larly, and laboriously masticated with salivary secre¬
tion, imparting to it the new and necessary quality
of ductility, and bringing about the other changes
already noticed.

: 5 8 BEES AND BEE-KEEPING.

Huber and Hunter both remarked that the common
idea, that wax had its origin in pollen, did not ap¬
pear to agree with observed facts— e.g., swarms
placed in empty hives carry little or no pollen, but
nevertheless build combs rapidly ; whilst the bees
of old hives, which construct no new cells, indus¬
triously carry home the many-coloured pellets.

Huber’s experiments,* intended to settle the question
of the origin of wax, are too important to be passed
over. He placed a swarm in a straw skep, and sup¬
plied it with honey and water, whilst so shutting in
the bees as to permit of full ventilation. The agita¬
tion of the captives passed away when their hive was
placed in a cool, dark spot. At the expiration of
five days, five white and very fragile combs had been
constructed. These were removed, and the experi¬
ment continued, as it might have been argued that
the pollen the bees contained at the beginning of
the trial had sufficed to yield the wax. After a
further imprisonment of three days, and feeding as
before, five other combs were formed. This proce¬
dure was repeated to the fifth time with similar results.
The experiment was now reversed, a swarm being
supplied with pollen, but not honey, and during
eight days of captivity neither wax scales nor cells
of comb were produced.

Huber had not failed to note that honey con¬
tained both minute quantities of pollen and, acci¬
dentally, scraps of wax, so his earlier experiment
was re-tried upon three swarms. The first received

“ Nouvelles Observations sur les Abeilles,” 1814.

WAX, AND BEE ARCHITECTURE.

!59

clarified sugar syrup, the second dark brown moist
sugar syrup, and the third honey. The results, as
narrated, were remarkable, and I hope hereafter to
test them, for Huber states that, uniformly, during
seven consecutive deprivations of comb, the wax
secreted by those fed upon honey was far less than
that yielded by those receiving sugar, of which the
dark brown gave invariably the highest quantities of
wax; but these were subsequently equalled by maple
sugar. It was thus established, that saccharine matter
from the nectaries of flowers, a:s honey, or in any
other form, was capable of .furnishing all the material
needed for the production of wax. But let us not
forget that comb building, even apart from the
salivary secretion needed to make the wax plastic
demands muscular and nervous wear, both occasioning
a loss of nitrogenous matter and salts—especially
phosphates—and these cannot be made up by sugar;
which, as a heat and force-former, contains, like wax,
only hydrogen, oxygen, and carbon. Physiology an d
prolonged experience alike, then, show that the effort
of_comb building is terrifically exhausting to the bee,
unless pollen or a substitute is at command, in
addition to sugar syrup, or even honey.

It is unusual,~aS—prffviously observed, to find bees
in the summer season without traces of wax in the
abdominal pockets, but these are frequently so thin
and impalpable that microscopic dissections alone will
reveal them. I received by post, on October 22, 1885,
a single bee, with a request that I would determine
its sex, as it was supposed to be a queen ! I found
it in all respects a genuine worker; it revived by

BEES AND BEE-KEEPING.

160

warmth, and during five days was regularly fed
on thinned honey.. Its liberal diet, aided by the cosy
solitude I gave it, enabled it to secrete wax, of
which I found, at . the “ post-mortem/' eight beauti¬
fully transparent scales. This little incident brings
before us the external conditions which aid wax
secretion.

When a swarm is placed in an empty hive, the
bees climb the sides, and gradually, and in close
order, advance along the roof, carefully securing
themselves by the hooks (anguiculi, page 124) of
the front legs, in order to sustain the weight of
lengthened chains of their comrades, formed by bee
after bee hooking her fore feet into the hind feet
of the one above. In this manner, the whole swarm
will in an hour or so suspend itself in festoons, which
are usually in part attached beneath to the neigh¬
bourhood of the hive door, in order that an efficient
guard may be kept up, and to give ready ladder-way
should any arrive with supplies. This arrangement
complete, all is hushed in perfect stillness, no bee
of the living chains moves, whilst a high temperature
is sustained ; and now the abundant food with which
each emigrant charged herself before she left the old
home comes under the process of conversion, and
the wax distils copiously on to the surface of the
thin membrane in the pockets. Wax is noT -ch&mi-
cally a fat or glyc eride, and those who have called
ilT^the fat "of bees' ” have grossly erred ; yet it is
nearly allied to the fats in atomic constitution, and
the physiological conditions favouring the formation
of one are curiously similar to those aiding in the

WAX, AND BEE ARCHITECTURE.

:6:

production of the other. We put our poultry up to
fat in confinement, with partial light, to secure bodily
inactivity, we keep warm and feed highly. Our bees,
under Nature’s teaching, put themselves up to yield
wax under conditions so parallel that the suitability
of the fatting-coop is vindicated.

The wax having been secreted, a single bee starts
the first comb, by attaching to the roof little masses
of the plastic material, into which her scales are con¬
verted, by prolonged chewing with secretion ; others
follow her example, and the processes of scooping
and thinning (presently to receive detailed attention)
commence, the parts removed being always added
to the edge of the work, so that, in the'darkness, and
between the bees, grows downwards that wonderful
combination of lightness and strength, grace and
utility, which has so long provoked the wonder and
awakened the speculation of the philosopher, the
naturalist, and the mathematician.

The comb (Figs. 3 and 4) is constructed on a middle
wall, or midrib (seen in the section at ab , A, Fig-
36), which forms the bases or ends of the layer of
cells (c, d) covering it on each side, and which are
hexagonal prisms, in length somewhat less than ^in.
The midrib B consists entirely of lozenges, or rhombs
(/.(?., figures with four equal sides and two acute
and two obtuse angles), of which each cell covers
three, constituting its base, as may be seen by the
double line representing the cross section of the cell
sides. The rhombs so meet, with an obtuse angle of
each in the middle of the cell bottom, that their edges
cannot be joined whilst they lie flat, as their enlarged

162

BEES AND BEE-KEEPING.

outline (r, r, r, C) in horizontal position shows. At B,
each three is thrown into a concave form. From this
it is evident that, if the cells on the two sides of the

Fig. 36.—Comb Structure.

A, Section of Comb (Natural Size)— ab, Midrib; c, d, Cells. B, Midrib, consisting
of Three Rhombs for each Cell (Natural Size). C, Rhombs and Cell Sides
Magnified Three times— r, Rhombs of Cell Base ; s, Sides; o, Obtuse Angles
of 109°; a, Acute Angles of 71°. D, Cardboard Pattern, which Folds into Two
Cells ; Lettering as Before. E, Rhomb Giving Ratios of Diagonals, ef and gh.
F, Cross Section of Comb, showing that the Inclination of the Rhomb does not
affect Storage Space. G, Section of honey- comb, Showing Curvature of Cells.

comb stood immediately opposite, the concave bases
of the one side would present the extreme incon-

WAX, AND BEE ARCHITECTURE.

venience of convex bases on the other, like, indeed,
the bottoms of those bottles which are made to look
large and hold little—the very opposite of a principal
requisite in comb structure; but equal concavities
are given on both faces, by the cell walls of one
surface coinciding with the adjacent edges of the
rhombs, which diverge from the centres of the cells
on the other (see A)—an arrangement easily under¬
stood by noting that the single lines dividing the
rhombs in B indicate the lines of the cell walls on
the remote side of the comb, while the double lines
indicate the cell walls on the near side. The same
fact may also be made apparent by piercing three
pinholes through the several rhombs of the base of
any cell, when these holes will be found to belong
to three different, though adjacent, cells of the
opposite face. Anyone really desirous of thoroughly
understanding this, and the other points yet to en¬
gage us, will do well to make, in cardboard, the form
given at D, where all the obtuse angles (marked 0 or
o'), and the acute (marked a), are equal to one another
respectively; the sides (s), if extended as far as the
edge of the letterpress, giving the correct proportions.
The dotted lines being half cut through, the form will
fold into two cells thrice natural size, and in correct
relative position on opposite sides of the comb, when
the edge x will fall on i', and the other numbered
edges meet as indicated. Designing a more com¬
plicated form, including two cells on each side, and
cut out in one piece, is an interesting, and not
excessively difficult, puzzle. Strips of gummed tissue
paper will hold the cells in form, which, when mdde

P 2

164

BEES AND BEE-KEEPING.

sufficiently large, I have found extremely useful for
lecturing purposes.

Supposing that comb equals its ideal or theoretical
form, Cramer’s* very elegant geometrical demonstra¬
tion shows that the angles of the rhomb must be
such that their two diagonals ( ef, gh, E, Fig. 36)
are to each other in the ratio of the side and diagonal
of a square; or, to use Cramer’s less popular, though
equivalent, form, the obtuse angle of the rhomb must
be such that its half has for its tangent 2. This
is only true of the angle 54 0 44' 8". The two angles
of the rhomb are, therefore, double the foregoing,
viz., 109° 28' 16", and its supplement, 70° 31' 44".
Thence, as geometric sequences, the angles at which
the sides of the prism (s, D) are cut at the base, in
order to fit on to the rhombs, is precisely equal to
those of the rhombs themselves ; and, further, the solid
angle formed at the apex of the pyramid, by the
meeting of the three obtuse angles ( 0, 0, 0, C) of
the rhombs will be equal to the solid angles formed
by the meeting of one obtuse angle (0) of the
rhomb, and the two similar obtuse angles (o', o', C)
of the sides. It is also true, that no other angles
give these equalities, which every geometrician will
recognise as affording the nearest approach to the
form of the larva possible to the number of plane
surfaces composing the cell.

It has sometimes been thought that these angles
gave greater space than any other; but this is an
error, as F will show ; for here the actual inclination

* Hutton’s “Mathematical Recreations,” or Huber’s “Nouvelles
Observations,” vol. ii., page 475.

WAX, AND BEE ARCHITECTURE. 165

gives no more room (if the material of the plate
be disregarded) than the dotted line ( hikl ), since just
so much as is taken out of the comb on the one
side is added to it on the other. The real economy
is in wax, for, had the midrib been flat, one-fiftieth
more of the precious secretion would have been

required; the midrib truly taking less, but the sides

much more, as the part now cut off from each of
them by the inclination of the rhomb must have

been added. The strength, also, would have been
diminished, while the shape would have been less
suitable for the accommodation of a round-ended
chrysalis. Maraldi, seeing the advantage of an

equality of the solid angles, such as previously
pointed out, calculated them upon the hypothesis that
they really were equal, making them iog° 28', and
70° 32', which is nearly accurate. To the same
author w r e are indebted for a comparison of the
results of theory with fact, by the admeasurement of
the actual angles of honeycomb. These he states to
be no° and 70°—as near an agreement as
could be expected. Out of the details now given,
by successive but individually small increments of
exaggeration, a most extraordinary myth has been
constructed, which, at last, asserts that Maraldi
submitted the problem of comb shape to Koenig,
and that his solution differed from Maraldi’s actual
measurements , made from comb , by only 2min. of a
degree (whereas Maraldi’s results were the outcome
of a geometrical hypothesis). The story continues,
that Koenig, being told of this discrepancy, and
examining his work for a third time, found an

i66

BEES AND BEE-KEEPING.

error in the logarithmic table he was using. Cor¬
recting his table, his results came into exact agree¬
ment with Maraldi’s measurements. Some, like
Lord Henry Brougham,* who shows much more of the
advocate than the philosopher, have, in consequence,
in a triumphant tone, asserted that bees have so
absolutely solved a most recondite mathematical
problem, that their work has actually corrected a
mathematician's press error. A story such as this,
once started, is certain of repetition, since, however
absurd, it has some sort of superficial prettiness, but,
like untruths generally, it degrades what it professes
to exalt; so let us examine its claim on our belief.
The difference of 2min. of a degree means a divergence
so small, that two lines forming this angle would
travel 144ft. before separating iin. from each other.
The length of the side of the rhomb being barely
-g-in., a divergence of 2min. on the whole length would
be about tt soq^ 11 -, an amount so small that a -yin.
objective would be required to give it visibility; but
the field of such an objective is about -^-in. dia-
meter, and in it not more than the tenth of each side
of the rhomb could be seen at once, upon which 2min.
would give only a distance which the mag¬

nificent quarters now produced, even under the most
favourable conditions, would be hopelessly unable to
resolve. The conditions, however, are most adverse,
while the comb, as a manufactured article, is ex¬
tremely rough, and, under a quarter, as irregular in
surface as the mud wall of barbarism. That Maraldi,
with the poor appliances of his day, did measure
* “Tracts, Mathematical and Scientific.” Griffin, Glasgow, 1866.

WAX. AND BEE ARCHITECTURE.

the angles of comb to minutes of a degree, needs
no contradiction.

The story is not distortion ; it is simple fable—a
fitting companion to “ Jack and the Bean Stalk,”
et hoc genus omne. But the whole thing is made
the more preposterous by the inexactitude of comb
itself. Careful measurements of the finest pieces
I have discovered, built with every advantage for
securing regularity, have shown that every cell is
far from geometrically accurate. It is difficult to find
a hexagon presenting errors of less than 3deg. or
4deg. in its angles, or, on an average, a distortion
more than a hundredfold as great as the 2min. in
question.

But because comb presents irregularities, must we
think less of it, or the little creature that moulds it,
or of the frame of nature of which the latter forms a
part ? Assuredly not; for if comb, to be perfect,
needed that kind of perfection which defective
reasoning, and an imperfect acquaintance with facts,
would have us to believe it to possess, then the incli¬
nation of the brood cells (Fig. 4), and curvature of
the honey cells (G, Fig. 36), suiting them so much
better to their purpose, would have been impossible;
and equally so, amidst many others, the fluctuations
in form to suit the character of the bee domicile, or
irregular transition cells to mingle drone comb with
worker. The instinct of the bee transcends the
mathematical solution that has been demanded in
achieving its true aim, which is economic. All
Nature, apart from the mystery of life, solves every¬
thing mathematically. The cricket ball flying from

BEES AND BEE-KEEPING.

the bat of the tyro, the spray from the maiden’s mop,
the tiny soap-bubbles of the laundress’s lather, as
much conform to perfect mathematical solution as
the path of a comet or the form of a star. One
November morning, about twenty years since, in my
early bee-keeping days, I found a skep turned over
on the ground, whither it had been knocked by the
scamper of a would-be burglar, who had to make his
escape before a vigilant representative of the law.
The bees, half benumbed, were crawling over their
combs, which showed but too plainly that they were
broken from their attachments. The difficulty was
beyond my powers. Now I should run a skewer
through the skep, and thread the combs upon it with¬
out removing them, but then I judged it best to lift
the combs as nearly as possible to the perpendicular,
put little wooden props between, place the floor¬
board over all, and turn to the erect position,
hoping for the best; but, alas ! the latter operation
was followed by a sound which filled me with dismay
—the combs had fallen ! I studiously fed, the bees
lived on, and, in the end, did well. But, by early
spring, their combs were a study. One was flat on
the bottom board, and was channelled beneath,

until it gave passage way in every direction. The
others, half down, were propped, and gnawed, and

repaired in such a way that their utility was not

much lessened; while, from the roof, new combs
were made to descend and join in sweeps into

their obliquity. I repeat, the mythic measurements
of Maraldi would degrade bee architecture. The very
atoms with which life deals yield mathematical results

WAX, AND BEE ARCHITECTURE.

169

always, but life so mingles and co-ordinates these
that the mathematics is masked, while her purposes
are secured.

Notwithstanding the absence of mathematical uni¬
formity in comb, it is manifestly a disposition of
parts of all others best calculated to afford a
maximum of strength with a minimum of labour, and
the greatest space for each cell, the quantity of
material being considered. On a plane surface,
where a number of small and similar spaces are
to be divided off by partitions, the hexagonal form
is the one which comprehends the largest area com¬
patible with the extent of the lines which inclose

Fig. 37.—Worker’s Jaw (Magnified Twenty-four times).
gf-rn, Great Flexor Muscle; a, Cutting Edge; sc, Wax Scales.

it; for the equilateral triangle, the square, and the
hexagon, are the only regular figures which admit of
being joined without interstices, and the proportion
of the area to the periphery of every regular polygon
increases as the figure consists of a greater number
of sides, and is, therefore, greater in the hexagon
than in either of the other two; besides, either a
triangular or square cell would form a most unsuit¬
able nest for a chrysalis with a round body.

But it is time that we endeavoured to understand
the manner in which the little artificer proceeds with
the wax which we have already seen attached to the

7 o

BEES AND BEE-KEEPING.

hive roof. She has jaws with a smooth edge ( a ,
Fig. 37), for scooping and moulding, and the closed
maxillae, with their polished surface, for a trowel.

As the burrowing wild bee chips out a hole circular
in cross section, to admit her body, so the wax-worker
carves into her wax, placing the material removed
upon the edge of the little pitting that increases
before her : but two points are accomplished of
which no good explanation can be given; first, that
the workers so place themselves that the concavity
made by one interferes with that made by her next
neighbour; and, second, that, when carving from both
sides, the scraping and thinning stops before an actual
hole is driven through. This mutual interference
forms into hexagons, cells that are always circular in
outline at the beginning. Let us try an experiment,
the apparatus for which is found in every home. A
floating soap-bubble is perfectly globular, because the
tension of the soap film covers the contained air by a
pellicle of the smallest possible area; but if we trans¬
fer the bubble to the surface of a saucer, its own
gravity flattens one side. Giving it now a companion,
the two will convert their films, where united, into a
perfectly flat wall, because the equal tension on its
two sides will throw the opposing curves into a path
between them. So two bees scooping in contiguous
cells, or one bee scooping alternately in two cells,
will, as the resultant of two opposite curves, produce
a straight side. Let us add to our two soap bubbles
five others, so that one occupies the centre, while six
surround it. Now, in cross section the central bubble
is perfectly hexagonal, while all contiguous walls are

WAX, AND BEE ARCHITECTURE. 171

flat, and those that are free curved, just as we dis¬
cover them to be in honeycomb, where every free
wall at the edge of the comb runs in a sweep, al¬
though partisans, like Lord Brougham, by example,
state the contrary. It has been advanced, in opposi¬
tion to this view of interference, that the outside cells
of the paper-nests of some wasps are angular; but,
as Darwin* hints, this is capable of explanation, and
I submit that it is clearly due to the necessary work¬
ing on both sides in alternation of three radiating
walls, and really lends confirmation to the position I
am arguing. To return to our bubbles. If a second
layer be placed over the first, not only will they be
hexagonal in cross section, but the superposed parts
of the two layers will frame themselves into rhombs
disposed in all respects like those of ideal comb. The
geometrical relations which embellish the wax tracery
of the bee are the necessary result of her mode of
proceeding. And mathematics is no more her endow¬
ment than it is that of the soap and water we have
been considering. These wonders come because the
whole creation is founded and sustained by the great
Geometer, whose laws of weight and measure neither
falter nor vary, so that, for the advantage of man,
the experience and observation of the past make him
the prophet of the future.

The costliness of wax to the bee, since it can only
be produced at the expense of many times its own
weight of honey or sugar, has led to great economy,
xlb. of it being moulded into 35,000 worker cells
in a case I carefully examined ; but an American
* “ Origin of Species,” chapter vii., “ Cell-making Instinct.”

BEES AND BEE-KEEPING.

writer states that he has noted 50,000 cells framed
from the same weight. The scraping is continued
until the walls are surprisingly thin ; those surround¬
ing the cells I never found thicker than -g-g^-in., while
some are only x^-in. The rhombs vary greatly, and
are stouter, reaching xg^-in. in some cases. Bees
will, under certain conditions, employ in comb build¬
ing shreds of wax which they have not secreted ;
and it is their habit to use up all nibblings and
scraps from neighbouring combs, so that a new
structure built between two old ones, containing
hatching brood, will be brown from the first, instead
of daintily white, the microscope showing it to
be not only full of the old cappings once lying over
the chrysalids, but to contain their cocoons, crossing
and recrossing in countless silken threads, while
pollen grains abound, a contamination from which
not even the cleanest super-comb is absolutely free.

The colour of a queen cell (A, B, Fig. 3) always
resembles that of the comb on which it is built, or by
which it is surrounded, because it is mainly made of
scraps, and for it little or no new wax is secreted.
Almost any material seems to be pressed into the
service, so that its great mass be made up, careful
searching generally being rewarded by finding, between
its layers, some of the cast skins of the contained
larva, which, though small, seem too useful to be
wasted. Brougham, having dissolved a queen cell in
“ terebinthine ” (turpentine), was sorely puzzled by
(“ Les Pellicules ”) the cast skins (see page 34), which
he did not understand, and for which he could not
account; but we must not dismiss the queen cell

WAX, AND BEE ARCHITECTURE. 173

without noticing its salient peculiarities. It is circular—
the typical form—in cross section, because it is built
alone, and is made to grow with the growth of the
grub it contains; and even if it have a companion
(for reasons given under Queen Raising), such cannot
be started so near that interference is possible ; and
as it is deprived of surrounding support, and exposed
to unusual strain—havitig to bear a cluster of bees
crowding round to give “ royal jelly,” and maintain
temperature—great strength is a necessity, and so
the economic labourers, that pare down worker cells
to the utmost limit, heap on material till it attains
forty or fifty times the thickness they ordinarily
allow. Yet their scooping instinct does not desert
them, as they pit the queen cell over every part of
its surface—an operation which saves material without
decreasing rigidity. But what is it that so perfectly
counterfeits mechanical wisdom, and prevents them
continuing this pitting to the limit reached in building
worker cells, which would inevitably wreck the nursing
cradle of their future queen, and so, perhaps, abso¬
lutely deprive them of all hope of a successor to a
lost mother ?

Liquid dyes kept within worker or drone cells for
weeks, have not, in any case, stained water lying in
the surrounding ones, which I have never found
other than perfect, notwithstanding the extreme thin¬
ness of the walls. The bees labour at both sides of the
latter, not only scraping the shreds, but rubbing them
into complete union with their maxillae, and this will
account for their freedom from faults; but observation
has led me to form a different opinion of the sealing

i 7 4

BEES AND BEE-KEEPING.

of honey-cells, which in former years I described as
air-tight. Most bee-keepers have noted that snow-
white sealed honey, if kept in a damp place, changes
colour, the sealing appearing to grow transparent,
and the honey itself not infrequently weeping. By
experiments and a microscopic examination, I have
made evident that former ideas were inaccurate, and
that not more than io per cent, at most of the sealing
of honey is absolutely impervious to air. To extract

a c B

Fig. 38.—Cappings op Cells, Various.

A Sealing of Brood Cell (Magnified Thirty-five times)— c, Cocoon ; w, Wall of Cell.
B Sealing of Honey Cell (Magnified Thirty-five times)—a, Sealing outside;
h,' Honey; w, Wall of Cell, Intervening Air left Black. C, Sealing of Brood
Deprived of Wax Shreds to show Cocoon, Debris, and Pollen Grains (Spot lens,
Magnified 200 times).

honey {see Extraction), it is necessary to shave off
the sealing; and if this be done skilfully, the wax
is removed so free from honey as to show at once
that the covers have never been in contact with the
cell contents. By consulting B (Fig. 38), we shall
see the reason of this. The horizontal position of
the cells prevents their being perfectly filled first
and covered afterwards ; but the bees, when the cell

WAX, AND BEE ARCHITECTURE.

is approaching fulness, cap its lower part, then add
honey, and increase the cover, placing shred upon
shred, after the manner a turf wall is built, until
the process is complete; no smoothing by the bur¬
nishing action of the maxillae on the inner side is
possible, and so the air (left black in the figure) inter¬
vening between the irregular tape-like shreds cannot
escape, and at the close forms a layer between the
honey and its cell-lid, giving increased whiteness
to the cover, and preventing also immediate leak, even
should a fault remain. The air being cut through
in uncapping, the caps are removed dry. Steeping
in water for three days a well-finished super contain¬
ing about 780 cells, all but forty-nine revealed that
they were defective, by losing their opaque whiteness;
for the honey had absorbed water, and was now in
contact with the inner wall. The practical import
of this observation will hereafter come before us;
but I must, at the moment, remark that the demand
for very thin capping, which one or two English
"judges” have made, is not wise, while the reason
they have given for preferring it is an error as to
fact.

Although the bee aims at compact coverings for her
honey, the sealing of her brood is made porous for
an object (as stated at pages 21 and 22), and, when
magnified in cross section (A, Fig. 38), shows the
looseness of its texture and the varied character of
its material, which is never white, and not even prin¬
cipally wax, only so much of the latter being used
as will bind the scraps and debris into oneness. On
the back, the cocoon threads ( c ) are seen catching

: 7 6

BEES AND BEE-KEEPING.

on to the prominences of the wax shreds or pollen
grain. One of these covers (from a drone cell by
preference), if washed in benzole, so as to dissolve
out the wax, and then mounted in the usual way
on a slip, forms a very interesting microscopic
object, especially for the spot lens, since this shows
the cocoon as bright golden threads on a black
ground (C).

The most puzzling of all variations remains to be
noticed, for no observer has discovered even the key
to the gauging of the dimensions of the cell by the
wax worker. It cannot be put in evidence that the
size of her body or head, or reach of her jaws,
determines it, for, under certain conditions which are
perfectly uniform, she discards the -g-in. diameter, and
starts constructing cells ^ of an inch between the
parallel sides, and these are used for the storing of
honey or the raising of drones, and so are commonly
called drone cells. The statement, many times made,
that twenty-five and sixteen of these respectively cover
a square inch is erroneous, as the outline is not
square, the correct numbers being as below:

—

Diameter.

length.

No. on Sq.

Side of

No. on Sq.
Ft. on One
Side of
Comb.

No. on Sq.
Ft. on Both
Sides of
Comb.

Worker cell ..

Drone cell .. ..

•Jin.

Jin.

-Lfm.

j^in.

4IS7

2660

8314
5320

The change

of size,

so mysterious

in its

cause,

cannot be made without disturbing the interfitting of
the hexagons, the difficulty being met by the con-

WAX, AND BEE ARCHITECTURE.

struction of so-called “ transition ” cells. The name is
misleading, and based on a misconception, for bees
pass at once from worker to drone, or vice versd,
and then build accommodation cells as necessity
determines, until the regularity of the new pattern
is established. It is singular that the form given to
these irregular cells, in all the books I have yet
seen, is such as no bee ever did or could con¬
struct, as it contains an acute angle bounded by
straight lines to the angular point. This matter is
not unimportant, for, if the books be believed in,

A, Impossible Cell—/, Angular Point 61° ; h. Head of Bee (Natural Size). B,
Comb, with Accommodation Cells—a, Normal Worker Cell; b , Pseudo-Cell;
c. Oval Cell; d, Normal Drone Cell; e. Truncated Angle, giving Room for
Bee’s Head.

the manner of cell elaboration cannot be understood.
Even Langstroth, to whom the debt of apiculture is
very great, has an illustration of the intermediate
cell with a prolonged internal angle of 62°, which a
number of English writers have improved (?) to 51 0 ,
whereas about ioo° is the limit the bee can reach. By
giving a copy (A, Fig. 39) of the cell (Fig. 48) of
Langstroth, into which I insert a bee’s head ( h), of
the natural size, the mistake becomes evident; for
how could this bee bring her jaws and maxillae into

BEES AND BEE-KEEPING.

the corner (f) 7 as, for reasons previously given, she
must, and that, too, from the very position in which
we have placed her, if the straight boundary lines
are to be modelled. The orthodox accommodation
cell, which , is really partially double, is seen
(Fig. 3) above two unsealed drone larvas, and
in it the septem is not continued to the top. A
few somewhat irregular forms, in addition to such
a one, will enable the bees to pass completely
from one size to the other. But even where the
greatest difficulties are presented, no angles of less
than ioo° are found— e.g., in B, Fig. 39, made from
a tracing of actual comb, constructed from pieces of
drone and worker, placed near to each other, for the
industrious little insects to repair and join ; a few
irregular cells are made to complete the accommoda¬
tion, the impossible angular point, of course, not trans¬
piring. The nearest approach to an angle lies at e,
where the width of the bee's head determines the
obtuseness, while at b the cell is only a depression,
not extending to the midrib, because its small size
prevents the entrance of the worker’s body. Such a
cell fills a gap, but is in no other way utilised.

It will be at once remarked, that the normal cells
in B stand between hexagons and circles. This is
true, more or less, of all comb, which, if cut through
in the middle of its cell partitions, shows these to be
very nearly straight up to the angles of the hexagons,
although some thickening is observable at the line of
junction ; but the end of the walls, at the face of the
cell, is always loaded by a rim of wax, which converts
the mouth into an approximate circle. This thicken-

WAX, AND BEE ARCHITECTURE.

179

ing of the rim exists in cells at all stages of their
progress, since the scraped-off wax is continually
added to the edge of the work. As this is reduced,
by being drawn out for lengthening the cell wall, it
is augmented by new supplies from the wax pockets
of the workers. The constancy of these thickenings
is essential to impart strength, permitting the clamber¬
ing throng to support themselves without fear of
breaking their own structure, which has its tenacity,
when completed, increased by being varnished with a
resinous body, called propolis, but whose qualities do
not yet come before us.

Fig. 40.—Details of Drone Cell (Magnified Twice).

A, Capping of Drone Cell, seen by Transmitted Light. B, Capping of Drone
Cell, seen by Reflected Light. C, Side View of Drone-Cell Capping (Section)—
a, Sealing Pervious to Air ; 6, Wax Struts.

The strain which the fragile-looking cells will bear
is extremely remarkable. One pound of wax, built into
35,000 cells, as before stated, will store 22lb. weight of
honey; from which it follows, that the wax of a cell at
the top of a comb, fully filled, and ift. deep, supports
22 x 60 = 1320 times its own weight. The special
manner in which the top cells are strengthened will be
most usefully considered under the head “ Foundation,”
in our Practical Section. But drone cells are less rigid

Q 2

:8o

BEES AND BEE-KEEPING.

than worker, in the ratio of 25 to 16; so that, in their
case, a system of girdering is adopted, which greatly
interested me when I discovered its existence a few
years since. If a sealed drone grub be dropped back¬
ward out of its cell, by cutting away the base of the
latter, the capping and its surroundings, as seen by
transmitted light, give the appearance of A, Fig. 40.
The porous and weak, but semi-opaque, sealing
occupies the centre, while the angles are made rigid
by filling up with transparent wax, which is done
with such regularity that an exceedingly pretty star-
like form results ; but the little engineers seem not
content with this provision, so they throw webs across
from the convexity of one cell to the convexity of the
next, each web radiating in six arms, as seen at B.
The utility of the arrangement as against downward
strain is evident, as the strap ( b , C) clearly prevents
any sinking. The illustration shows the almost
hemispherical form of the c'ap, which, we must
remember, is made by the bee outside the cell, so
that the convex side is towards her. How this form
is accomplished I know not, and my difficulty is but
increased by learning that the contained grub gives
no help by its presence within. While studying the
drone, I cut, as I imagined, about a hundred of their
sealed larvae from a hive, for dissection purposes. The
cappings were fully as prominent as usual, but, to my
astonishment, I found worker larvae within, and these
only; and searching further revealed the curious fact,
that the queen seemed incapable of laying a drone
egg, of which more hereafter. The evidence of
interference, giving form to worker cells is so con-

WAX, AND BEE ARCHITECTURE.

181

elusive to the unbiassed mind, that I cannot help
supposing that it possibly applies in this matter. If
I ventured on a theory, it would take this form. The
throwing up of the cell walls from the three contiguous
obtuse angles of three adjoining cells produces the
triangular piece (transparent in A), with a depression
in its centre, and which an examination of every
drone comb (store or brood) will reveal. The worker
pitting these concavities forms the wax pieces which
strengthen the angles, as previously mentioned,
while their edges, naturally becoming prominent,
furnish the six straps, by simple junction, when,
between the latter, a comparatively flat sealing is
thrown across.

Pure wax is perfectly white; the propolis added
as a varnish is the usual, though by no means invari¬
able, source of its yellow colour, which may depend
upon some peculiarity in the nectar the bee is gather¬
ing at the time of building; but combs in which breed¬
ing has taken place are always more or less brown.
This has been explained by stating that the cast
skin of the grub causes the discoloration. The cast
skin, however, is a delicate and transparent pellicle,
and gives no colour to the comb. We have already
learned that the toning is due to the residua of the
bowels, plastered outside the exuvium, within the cell
wall. This material at first fills the corners, as may
be seen by examining cells in which one hatch only
has occurred, when the angles will be dark, while
the sides will be only very slightly stained. In this
connection, Fig. 4 may be examined with advantage.
After a few hatches, all angularity at the cell base

BEES AND BEE-KEEPING.

will have vanished, and the cross section will be
nearly circular—the typical form, again, to which the
cells in this way are always approximating.

The details that have passed before us, and of which
hereafter we shall see the practical import, are
many and various. Are any disappointed that, during
their discussion we have deprived our bee of the
mathematical laurels some would force upon her little
brow? It cannot be; for surely we have not dis¬
graced her. Rather have we given her new honours, by
disclosing adaptations and variations truly astonishing;
while all that we have said has not removed her from
the front rank of dumb artificers : for even man, with all
his art, has not been able to give to wax equal beauty
to that it yields at once to the simple tools of its
own producer. Yet that which is brick, mortar, and
wood, to the bee, must mainly strike us in its
utilitarian aspect; for her combs are rows of rooms
unsurpassably suitable for feeding and nurturing the
larvae, for giving safety and seclusion during the
mystic sleep of pupahood, for ensconcing the weary
worker seeking rest, and for safely warehousing the
provisions ever needed by the numerous family, and
by all during winter’s siege. Corridors run between,
giving sufficient space for the more extensive quarters
of the prospective mother, and affording every facility
to the busy throng walking on the ladders the edges
of their apartments supply ; while the planning of
the whole is such that the exactions of modern
hygiene are fully met by air, in its native purity,
sweeping past the doorway of every inhabitant of the
insect city.

CHAPTER XII.

STING STRUCTURE.

Stings and Ovipositors—The Piercing Apparatus —
Sheath, Darts, and Barbs—Groove and Slide Rod —
Muscular System—Method of Making the Wound
—Nerve Structure—Poison Gland—Formic Acid —
Experiment — Poison Bag — Pump-like A ction —
Removal of Sting—Sting Palpi—Lubricating
Gland : Its Secretion ; Microscopic Examination —
How Friction is Reduced—Queen Sting: Its Size
and Modifications; How Planned for Removal;
Why Curved; Often Atrophied; Development;
Necessity for.

But few of those interested in these pages have
not, in times gone by, tasted of the potency of
the instrument now to occupy us, and the remem¬
brance of its stimulating efficacy, apparently so out
of proportion to its size, may quicken our interest
as we investigate its structure, which we shall find
as complex and remarkable, and equally as suited to
its purpose, as those that have already come before us.

Amongst bees, as well as the other aculeate
Hymenoptera, the sting is exclusively the endowment
of the females, and while its primary use is to

184

BEES AND BEE-KEEPING.

arm its possessor, it is also probably helpful in the
deposition of eggs. Anatomically, it is analogous to
the boring ovipositor of the saw, gall, and ichneumon
flies, insects belonging to the same order as the
bee. Whilst the ichneumon deposits her eggs in the
soft bodies of other insects (which must, of course,
first be pierced by her sharp ovipositor), the saw
and gall flies have really to cut, by means of rasping
teeth, an aperture into leaves, buds, or even timber,
so that the eggs may be inserted, together with a
droplet of fluid which has a peculiarly irritating
effect upon the vegetable tissues, occasioning the pro¬
duction of the galls, which are new growths, that
serve not only to protect the larvae the eggs furnish,
but also to afford them nutriment. When we call to
mind the strange piercing power of the sting, and its
venomous effect, we shall have no difficulty in accept¬
ing the statement that the difference between the sting
and the ovipositor is rather that of function than struc¬
ture ; they are both situated at the posterior region of
the abdominal cavity, the latter being usually carried
in a prominent position, whilst the former is always
hidden when in repose. Let us now consider the
mechanism by which the worker bee forms the wound
when she strikes.

The piercing apparatus consists of three main
portions—a so-called sheath and two darts. The
former ( sh, A, Plate VI., and side view, E, Plate VII.)
is a dark brown and strong chitinous piece, large and
pouch-like at the upper, but narrowed and flattened
considerably at the lower part, where it terminates
in an extremely thin cutting edge, which is the

STING STRUCTURE.

first to enter when the sting is used. The puncture
having been made, the sheath is held in the wound
by two* rows, each containing three, or unusually
six, microscopic teeth (E, Plate VII.), pointing back¬
wards, and acting like the barbs of an arrow or
harpoon. The sheath has three functions: first, to
open the wound as we have seen; secondly, to act
as an intermediate conduit for the venom ; and,
thirdly, to hold in accurate position the long darts ter¬
minated by barbs ( b, A, Plate VI., and E, Plate VII.).
The sheath, so-called, does not inclose the darts
as a scabbard, but is cleft down the side presented
to us in Plate VI., which is below when the sting
points backwards. This cleft at the upper part
of the sheath, where the latter is oval in cross
section, is just wide enough to permit the two darts
to close it by standing side by side between its
edges. But as the darts move up and down at their
pain-inflicting work, they would immediately slip
from their position, unless prevented by a mechanical
device, exhibited by B and C, Plate VI., giving
in cross section sheath and darts near the termina¬
tion, and at the middle of the former. The darts
{d) are each grooved through their entire length,
while upon the sheath (sh) are fixed two guide rails,
each like a prolonged dovetail, which, fitted into the
groove, permits of no other movement than that
directly up and down, to which we have previously
referred. At E, Plate VII., the dart has been

* Mr. Hyatt, who has carefully examined the sting [American-
Quarterly Microscopical Journal, vol. i.), has found only one row, but
two always exist, although they are difficult to bring into view.

i86

BEES AND BEE-KEEPING.

forcibly dragged from its position up to S. The
darts are terminated by ten barbs, of ugly form (D,
Plate VI.), and much larger than those of the sheath,
and, so soon as the latter has established a hold, first
one dart, and then the other are driven forward by
successive blows. These, in turn, are followed by
the sheath, when the darts again more deeply
plunge, until the murderous little tool is buried to
the hilt. But these movements are the result of a
muscular apparatus yet to be examined, and which
has been dissected away to bring the rigid pieces
into view. The dovetail guides of the sheath are
continued far above its bulbous portion, as we see by
E, Plate VII. ; and, along with these, the darts are
also prolonged upwards, still held to the guides by
the grooved arrangement before explained ; but both
guides and darts, in the upper part of their length,
curve from each other, somewhat like the arms
of a Y, to the points c, c (A, Plate VI.), where the
darts make attachment to two levers ( i, i'). The
levers ( k , /, and k ', l') are provided with broad
muscles, which terminate by attachment to the lower
segments of the abdomen. These, by contraction,
revolve the levers aforesaid round the points f, f, so
that, without relative movement of rod and groove,
the points c, c approach each other. The arms of
the Y straighten and shorten, so that the sheath and
darts are driven from their hiding place together,
and the thrust is made by which the sheath produces
its incision and fixture. The sides being symmetrical,
we may, for simplicity sake, concentrate our attention
on one, say the left in the Figure. A muscular con-

STING STRUCTURE.

traction of a broad strap joining k and d (the dart
protractor) now revolves k on /, so that a is raised,
by which clearly c is made to approach d — i.e., the
dart is sent forward, so that the barbs extend beyond
the sheath and deepen the puncture. The other dart,
and then the sheath, follow, in a sequence already
explained, and which G, Plate VII., is intended to
make intelligible, a representing the entrance of the
sheath, b the advance of the barbs, and c the sheath
in its second position. The barb retractor muscle
is attached to the outer side of t, and by it a is
depressed, and the barbs lifted. These movements,
following one another with remarkable rapidity, are
entirely reflex, and may be continued long after the
sting has been torn, as is usual, from the insect.
By taking a piece of wash-leather, placing it over
the end of the finger, and applying to it a bee held
by the wings, we may get the fullest opportunity
of observing the sting movements, which the micro¬
scope will show to be kept up by continued impulses
from the fifth abdominal ganglion, and its multitu¬
dinous nerves ( n , A, Plate VI.), which penetrate
every part of the sting mechanism, and may be
even traced into the darts. These facts, together with
the explanation at page 49, will show why an ab¬
domen separated many hours may be able to sting
severely, as I have more than once experienced.

But it is not the laceration from the sheath, nor
the punctured wound of the dart, that makes the
insect robber of honey so cautious, nor man so
solicitous to conciliate the gracious favour of Miss
Apis; for, when the worst has been done, we have

BEES AND BEE-KEEPING.

a wound whose maximum depth is xg-in., and whose
diameter is -g-^in., or less than of the area of
that inflicted by a common pin. The sting derives its
value, as an instrument of attack and defence, from
the poison with which it is associated, and which is
derived from a gland ( pg , A, Plate VI.) having often the
astonishing length of i-^in. in the worker and ijtin.
in the queen. In the former, dissection reveals it tra¬
velling, like two attenuated, nearly transparent threads,
over the outside of the chyle stomach, while its ends
are swollen into forms resembling the plumber’s iron.
These are full of curious dotted cells, containing
granular matter, and are abundantly supplied with
tracheae. The bifid gland unites at some distance
from the poison sac ( ps ). Its structure (H, Plate VII.)
is in the divided portion intracellular, and the duct-
lets of the cells (I) may be brought into view by treat¬
ment with liquor potassa.

The poison it secretes is formed from the blood by
cell-elaboration and transformation, and its active
principle seems to be formic acid, probably associated
with some other toxic agent. If a bee be made to
sting a piece of paper stained with litmus, which
is a common test for acids, the dye is immediately
reddened. On this account, ammonia is often recom¬
mended to allay the irritation a sting causes, as it
is argued that the alkali must act as a neutraliser.
Formic acid is poison to the blood of the bee, which
dies by a sting from its relative, although it is not in¬
jurious if taken, in reasonable doses, into the creature’s
stomach, as food mixed with it is accepted readily, while
no untoward consequences are observed (see Diseases).

STING STRUCTURE.

The poison bag, of considerable size, is lined with
epithelium, but is not muscular, as stated by
Mr. Hyatt, its venom being driven from it by the
play of the muscles giving activity to the' apparatus,
and by a singular pump-like arrangement, presently
to be noticed. Indeed, a comparison of the highly
muscular poison sac of the wasp, which has no
valvular appendages, with that of the bee, is highly
instructive. The poison sac contracts into a strong
neck ( pb , E, Plate VII.), which enters the upper
bulbous part of the sheath, in which play the valves
( va ). The walls of the sheath are double, with blood
between, while it is into the cavity within the in¬
terior lining that the poison enters, and so bathes
the back part of the surface of the darts, which
stand in the cleft in the front of the sheath, as
before stated. Since the darts present concave sur¬
faces to each other, they inclose a tube-like space
(p, B, Plate VI.) between them, through which the
poison passes downwards, towards the base of the
wound, being driven forwards by the piston-like
action of the valves (va, E, Plate VII.), which
descend with the stroke of the darts to which they
are fixed, and sweep the poison before them, ram¬
ming it onwards, when the end of the stroke is
reached, by the valve meeting the lower end of the
pouch. These valves are remarkable structures, and,
with regard to them, I venture to differ absolutely from
Mr. Hyatt. It will be remembered that the bulbous
part of the sheath is oval in cross section. The
greater diameter of the oval runs from front to back,
and this space the two valves divide between them,

BEES AND BEE-KEEPING.

190

one taking the right, the other the left, half.
As they pass up and down in company with the
darts, they never become absolutely clear of each
other. Although one be at the top, and the other
at the bottom of the stroke, they still, in part, are
side by side, so that clashing or interference is im¬
possible. Each is formed from the dart, by the
throwing back of two strong, parallel, chitinous, rod¬
like pieces, supported by a truss {tr, L, Plate VII.)
above them. As the whole valve is necessarily
narrow, the space between these rods is small.
Above the truss we find a feathery expansion, in
the form of a hood, which really holds in position a
most delicate membrane — not represented in the
Figure—really a bag, mouth downwards, the edges of
its mouth being attached to the parallel rods, so
that, when the down stroke is made, the poison, with
which the pouch is always full, passes into, and
expands, the bag, as a butterfly net is opened out
when it sweeps through the air. Below each rod
depends another membrane, semicircular in outline,
and stiffened by numerous chitinous, branched thicken¬
ings, seen above va. These flaps, at the down
stroke, separate from each other, and the better
drive the venom before the advancing valve. At the
end of the stroke, the fully-extended membranous
bag, by its elasticity, continues to drive on the
poison until its companion takes its place. At the
upward stroke, the bag collapses, and settles on to
its feathery support, which holds it in position for re¬
filling, while the depending flaps fall together. The
accurate fit of the darts prevents the escape between

STING STRUCTURE.

191

them of the poison, which is constantly being
pumped forwards, as we have seen ; but exit is pro¬
vided by minute channels ( 0 , 0, o , 0 , D), passing
from the poison cavity to the base of each of the
five lower barbs ; the poison is thus sunk to the
lowest point in the wound, where it collects, so
long as the sting remains, until the poison sac is
itself empty. From all that has been said, it is
apparent that the more quickly the attached sting
can be removed, the better. A prompt brushing
of the finger over it, or rubbing of the hand, if
wounded, rapidly over some part of the clothing
before more than a superficial puncture has been
made, will usually dislodge the entering sheath ; but
even if the sheath and darts have descended their full
length into the skin, every additional thrust, although
adding nothing to the depth of the wound, still
pumps into it additional virus.

The bee, quick as thought in the execution of her
attack, nevertheless does not inflict a wound until she
has examined the nature of the surface to be punc¬
tured, using a pair of very beautiful organs {p, p , A,
Plate VI.), called palpi, elaborately provided with
feeling hairs and thin nerve ends. She is never so mad
with anger but that she has method in her madness,
preferring animal to vegetable substances for attack.
It is extremely difficult to get her to sting writing
paper, and some substances (to be mentioned under
Practical Management), applied to the skin, will almost,
if not absolutely, save it from attack.

The strictly mechanical build of the sheath and
darts—reminding one almost of the guide rods of a

ig2 BEES AND BEE-KEEPING.

steam engine—introduces a question which greatly
puzzled me before I found its solution. If some of
the virus, exhibited as a tiny drop at the point of
the extended sting of an angry worker, be removed
by a glass slip, and allowed to dry for three or four
minutes, it will become hard, leaving a little promi¬
nence, as though it had been gum water ; and if it
be placed under the microscope as it sets, it will be
seen to split, by contraction, into lines, which rapidly
travel across the field of view. Dr. Bevan* says :
“ If the poison be looked at by a microscope, pointed
crystals will become visible. These may be seen at
first floating in the venom, and gradually shooting into
crystals as the fluid part evaporates.” Careful experi¬
ment proves that Dr. Bevan was probably deceived by
a defective microscope. He mistook, no doubt, the
fissures for crystals. The object is a, curious one, and
the experiment so easy that it should be tried. But
to our point* How is it that this gummy body, insinu¬
ating itself between the grooves and tenons, does not
quickly fix them together, and render the sting utterly
inoperative ? Another gland, not seen in the Figure,
prevents what some might consider “ a consumma¬
tion devoutly to be wished.” Its place is behind
the ganglion ; it is much smaller than the poison
gland, being about ^-in. long and y^-in. in dia¬
meter, and, like its companion, it enters a sac,
which is the reservoir of its secretion, and which
would be situated behind the vulva ( v ) in the Figure.
The fluid it produces is a lubricating oily body, which,
entering between the working parts, secures their

* Bevan “ On the Honey Bee,’’ 1838.

STING STRUCTURE.

free play upon each other, while the sting itself
has as little contact with the venom as a duck’s
back with water. The extrusion of the sting brings
forward this secretion, which emits the peculiar odour
sometimes to be recognised if a number of bees are
roused to anger. When the poison is examined
through a good objective, the tiny oil globules which
have been provided by the lubricating gland are found
in thousands. Not only is the clogging of the moving
parts prevented by this beautiful system of lubrica¬
tion, but that friction is greatly reduced which
tells so terribly against long rods moving in grooves,
especially such as these* only g - 0 * 00 in. in diameter,
and fully 600 times as long as broad. And here I find
the very highest degree of mechanical perfection
is reached, by not permitting the rod to move in
absolute contact with its groove throughout its whole
length; for if the rods, at their upper parts (near
c, c', A, Plate VI.), be torn from their places, they
will be found to carry studs, or cogs, at regular
intervals, which themselves only come against the
back and sides of the grooves, so that they not only
diminish the contact surfaces, but act as distributors
of the lubricant—an antitype of the plan often
followed in machinery required to act with great
smoothness and precision. It must, however, here
be noted, that good high power objectives are re¬
quired.

The sting of the queen differs from the worker’s
in many particulars, although the plan of the struc¬
ture of both is identical. The worker uses her
weapon at great risk to herself, for frequently, and,

i 94

BEES AND BEE-KEEPING.

indeed, generally, she loses, not only the sting and the
venom gland and sac, but also the lower portion of
the bowel, so that her death follows in an hour or
two. The queen, whose individual life is bound up
with the very existence of the colony, carries a sting
which her instinct forbids her to use, except possibly
in the sole case of contest with another queen. She
may, by violent usage, be induced to protrude the
weapon of offence, but never does she in human
hands inflict a puncture. The instrument she carries
is also especially planned to prevent the catastrophe
which so frequently follows its use in the case of
the worker, while she receives from it superior protec¬
tive power because of its larger calibre and greater
length, the sheath being able to penetrate xg-in., and
the darts ^gin. more, making together yyin., while
the darts are -j^-in. in diameter. The sheath is more
heavily barbed than the worker’s, carrying two rows of
retrorse teeth, five or six in a row ; but the darts are
occasionally plain, though more often provided with
three minute teeth, which scarcely rise above the
general surface. It will be remembered, that the
venom escaped from the worker’s sting by tiny holes
beneath the lower barbs of the darts. Since the
queen’s sting is here practically barbless, exit for the
poison is given by hollowing out the inner faces of
the extremity of each dart into the form of a gouge.
When a worker stings, and becomes—as a friend
observed—quite unpleasantly attached to us, it will,
if allowed time, generally carry its sting away by
travelling round upon the wound, giving the instru¬
ment a screw movement, until it is free. The queen

STING STRUCTURE.

l 95

has been known, when stinging a rival, to so free
herself; and the form of the sheath presents every
opportunity, in her case, of securing this desirable
object. Its flatness and extreme hardness—for it
turns the edge of the finest razor—causes it to act
as a drill, so that, after a few turns, a large hole
is made, and it is clear, for, when the sheath is
freed, the darts offer no impediment.

It has been remarked, that the decided curvature
of the queen’s sting (q, F, Plate VII.), in contrast
to the straightness of that of the worker ( w ), is in¬
tended to give her such an advantage in combat, that,
while her sting is applied, her antagonist should be
powerless to reach her, so that a queen duel may
not be fatal to both ; but the curvature appears to me
rather to refer to mating and ovipositing, as the ex¬
tremity of the sheath can be turned far more com¬
pletely out of the way through its deviation from the
straight line, and the more so because the terminal
ventral plate ( r ') is much truncated, so as to afford
a recess into which it can be dropped. It cannot
be doubted, that the possession of the sting by
the mother-bee of the hive, at the same time that
it is generally denied to all but aborted females
(neuters), indicates that it has only a relation to
some special phase of bee-life, which observation
proves to generally transpire before impregnation ;
and it is curious that, in the great number of queens
I have dissected, a marked majority have had the
poison gland atrophied, while the poison sac, although
distended, has contained only a yellowish substance
almost, if not quite, as solid as new putty, and which,

ig6

BEES AND BEE-KEEPING.

of course, could not have supplied anything to a
wound the sting might have produced. Coupling
this with the absence of a special ganglion, such as
we find in the worker—for the last ganglion is mainly
required for the reproductive organs—the very secon¬
dary importance of the sting to the queen can hardly
be questioned.

The development of the sting during the larval
and chrysalis conditions is extremely interesting. Its
first indication consists in line prominences, or warts,
found in pittings on the ventral side of the penulti¬
mate and anti-penultimate segments of the maturing
larva [b, c, A, Fig. 47) ; but these are quite invisible
until, by hardening with alcohol, they make their
appearance beneath the external skin. These in¬
crease, and gradually assume the mature form during
the chrysalis condition, at the same time that the
segments bearing them diminish, especially on the
ventral side ( b , c, B) ; ’ so that, although appearing at
first on two distinct segments, the parts get fused
together, and the last segment but three (d, A) of
the larva becomes the last of all in the bee (d, C),
the intervening ones being introverted. The residue
of the disappearing segments is, at the same time,
modelled into the various parts that are accessory
to the complex organ, which, from the very manner
of its formation, lies within the body, like a sock
which has the foot turned inside the leg.

However much we may regard the possession of a
sting by a domesticated creature as undesirable, there
is no room to doubt its necessity to the bee in a
state of nature, where, in its hollow tree, or recess

STING STRUCTURE.

x 97

in a rock, the avenues are wide which would give
entrance to the robber. Even with the narrow door¬
way of a hive, bees are sometimes sorely worried in
the fall by the persistent attacks of hungry wasps,
that would overmaster any number of brave defenders,
were the latter deprived of their poisoned darts.
Man, by observation, and a knowledge of the habits
of the insect, can nearly always successfully prevent
or evade her attack; for it is too much to expect her
to concede that the master robs by right divine, or to
understand that he but levies a righteous tax upon the
prosperity he brings by the refinements of civilization
and the wisdom of his government—a failure in which
she has been followed by some higher in the scale
of creation than herself. There is, besides, a charm
in overcoming difficulties. Man was born to conquer;
he was placed in the world to “subdue it’’; and so
the zest of successfully marshalling, at our will, a
throng that could, if they knew their power, drive
us writhing from their neighbourhood, is far greater
(even though the profit might be less) than could
come, in the absence of the sting, from the man¬
agement of—

“ A golden hive, on a golden bank,

Where golden bees, by alchemical prank,

Gather gold instead of honey.”

Man and bees alike live in a world where good and
evil grow together, where the thrift of the industrious
excites the cupidity of the idle, where meurn and
tuurn are regarded sometimes as convertible. Let us,
then, accepting the sting without regret, strive to learn
the way in which, for us, it shall cease to be an evil.

CHAPTER XIII.

ORGANS OF THE DRONE.

Function of the Drone — Description of Organs —
Production of Spermatozoa—Development of Drone
after Imago Condition is Assumed — Microscopic
Mounting and Staining — Spermatophore — A rmor
Copulatrix—Eversion by Finger Pressure—Glove
Experiment — Extrusion of Spermatophore: how
Accomplished—Fertilisation in Confinement a
Failure — In-breeding Prevented — Survival of
the Fittest — Death of the Drone — The Queen
becomes Double Sexed — Parthenogenesis — Herm¬
aphrodite Bees : Reason why so many are pro¬
duced—Natural Selection—Drone Slaughter.
BEFORE entering upon a detailed examination of the
drone (Fig. 5), we should know something of the posi¬
tion he has to occupy. The queen—whose egg-laying
powers have already come before us—on the wing, and
when a few days old, mates, but never again, however
much her life may be protracted. The drone at this
time gives a sufficient amount of fertilising material
to secure the individual impregnation of the multi¬
tude of eggs afterwards to be laid. To enable him
to accomplish this, his organs are, in some respects,

ORGANS OP' THE DRONE.

199

unique, and of most disproportionate size, greatly ex¬
ceeding those of even the largest Bombus , for the

Fig. 41.— Organs of Drone (Magnified Twelve times).

A, Organs Removed from Body, but in True Relative Position—t, Testes ; vd, Vas
Deferens; vs, Vesicula Seminalis; mg, Mucous Glands; de. Ductus Bjacula-
torius; 0, Termination of Organ; s, Sickle-shaped Scale, beneath which
Spermatophore is formed ; ts, Triangular Scale ; b. Bean; /, Fan-shaped
Appendage ; r, Ridges (Five-banded Piece of Swammerdam); h, Horns; m.
Masque of Reaumur, or Hairy Membrane. B, Spermatozoa Developing within
Spermatic Tube of Testes (Magnified 500 times)— sv, Spermatic Vesicle; n.
Nerve Cells. C, Spermatozoa as they arrange themselves after Removal from
the Body—a, Coiled Form ; h, Head ; th. Thread. D, Face View of Appendage
/ in A—/", Fan-like Fringe. E, Organs Extruded ; lettering as A. F, Front
View of Portion of Bean— s", Sickle-shaped Scale; ,sp, Spermatophore; ts",
Triangular Scale.

200

BEES AND BEE-KEEPING.

e gg-p r °ducing capabilities in this genus are relatively
restricted. Since he has but one function, he is
needed only during the months when swarming may
be possible, so that, normally, in the winter he is non¬
existent.

The distinctive sexual organs consist of a pair of
testes ( t , A, Fig. 41), communicating, by narrow tubes
(the vasa deferentia, vd), with the vesiculse seminales
(vs), which discharge, by their small extremities, into
the large mucous glands (mg), at whose junction
originates the ductus ejaculatorius (de), terminating
at 0 —the beginning of the true organ of generation,
which, in the condition of repose, lies outside in, and
so within the cavity of the body ; while, in activity, it
assumes the form given at E.

The testes are tender, white bodies, slightly flattened,
and much smaller in the adult drone than the ovaries
of the queen, to which they are homologous— i.e.,
they are to the drone what the ovaries are to the
queen. They lie within the abdomen, at its upper
part, and on each side of the digestive organs. Sper¬
matic tubes, or canals, to the number of about 300,
which open upon the vas deferens, make up nearly
the whole of their substance.

In the male chrysalis the testes are already not
only existent, but of enormous size, equalling at this
time, perhaps, the ovaries of the queen. They lie over
the then blind intestine, toward the dorsal surface.
Their canals are, at this time, filled with spermatic
vesicles (sv, B, Fig. 41), and with filamentous sper¬
matozoa, many of which are endowed with a lively
movement. (We must somewhat anticipate, by saying

ORGANS OF THE DRONE.

201

that these active threads are, really, the instruments
of impregnation, and have to be transferred to the
female.) As they lie, by tens of thousands, in parallel
lines, undulating rhythmically through their whole
length, they have, under the microscope, a most extra¬
ordinary appearance, which has been likened to a field
of barley oscillating under a gentle breeze. When the
drone quits the cell, the testes are still very large,
and extremely active, and but few spermatozoa h a^e
been transferred to that part of the body from which
they can be discharged, so that th e drone is not
at this time, nor is he for several subsequent days,
fully fit to accomplish the purpose of his existence.
But the virile threads, maturing rapidly, keep passing
from the testes to the vesiculae seminales, which
become now completely charged with them. It is
by opening the vesiculae at this stage that spermatozoa
are obtained in the best condition for the microscope.
They are such wondrous objects that I will explain
a method I have pursued most successfully in mount¬
ing them. Secure a drone (not newly-hatched), as he
is perambulating the combs, open the body, remove
the vesicula, break one end, and, with the forceps,
apply, for a moment, the ruptured part to the sur¬
face of some glass covers upon which a small
quantity of water has been placed (one vesicula will
give a supply for a dozen slides) ; leave to dry,
keeping from dust; warm in the flame of a spirit
lamp to set the albumen, pour on each three or
four drops of watery solution of Spider’s purple, and,
after five minutes, wash, dry, and mount in Canada
balsam. For critical examination with high powers,

202

BEES AND BEE-KEEPING.

spermatozoa should be mounted in glycerin. If stain¬
ing be desired, a minute quantity of the purple
added to the glycerin will accomplish it, as, in a
few weeks, the spermatozoa will have absorbed every
trace of the dye.

To return. The testes, although retaining partial
activity, shrink and flatten as the drone reaches virile
maturity. The mucous glands, secreting a slimy
liquid, give to the separate spermatozoa some cohesive
power, presently utilised. The spermatozoa, mingled
with mucus, pass continually onwards, through the
ductus ejaculatorius (de), into the bean, where a
mysterious arrangement of the myriad threads occurs.
They fill up the bean (b), and their mass is now
denominated the spermatophore (seen lying under ts
in the Figure). The ductus ejaculatorius has walls
of great muscularity, and, in the act of mating, it is
one of the main forces for putting the organ right
side out, so that it becomes external to the body.
The rounded little white, and somewhat fleshy, part,
the bean ( b ), is united to two brown, crescent-shaped
scales (j), and two triagular ones (is), which are rudi¬
ments of the usual armor copulatrix of the Hymeno-
ptera. The bean, and the remaining parts, from o
to m, are surrounded by a membranous sheath, which
remains intact after the expulsion mentioned above.
The curious, bright brown ridges (r, A, and r', E) hinder
the withdrawing of the organ during coition, and aid in
tearing it, according to rule, from the body of the male.
Below the ridges are found two membranous sacs
(, h ), which are always more or less filled with air, and
have been called pneumophyses from this fact. In

ORGANS OF THE DRONE.

203

repose these parts are bent and flattened, but when
swollen they become hard and resistent, and take
the form of divergent horns, which, as they expand
in undergoing eversion, pass into and fill the bursa
copulatrix of the queen—a recess on both sides of
the vagina. If a drone (by preference, one caught
as he settles on the alighting-board, from his quest
on a fine summer day) be pressed between the
fingers, from the thorax towards the end of the
abdomen, his organs will be expelled, because the
- air sacs are full, and the whole animal in an excited
condition. They will then, wholly or in part, assume
the form shown at E, and may be dried, attached to the
body, and kept for subsequent study. This experi¬
ment will enable us to understand the manner and
order of the extraordinary changes that occur during
coition. Under excitement, indeed, the extrusion fre¬
quently happens to the drone without any mechanical
squeezing, a sudden decapitation not unusually occa¬
sioning it; and dead drones, with the organs ex¬
ternal, may constantly be found about the apiary in
the summer.

The spermatophore, ovoid in form, gives, by in¬
terior pressure, a bulbous shape to the upper part of
the organ, which lies loosely within its sheath in the
cavity of the abdomen, and adherent to the body at
the edges of the sexual orifice only. If a glove
have one of its fingers turned outside in, the latter
being then filled with some semi-fluid matter, such
as paste, it may illustrate the action at the time of
the accouplement. The junction of the finger with
the hand of the glove will represent m, A (Fig. 41),

204

BEES AND BEE-KEEPING.

the top of the finger the termination of the organ,
o, while the hand will be equivalent to the abdomen.
If we blow violently into the glove, the finger will
be extruded. But, to complete the illustration, the
top of the finger should have an aperture similar to
that at o, by which the spermatozoa composing the
spermatophore enter the bean. Repeating our experi¬
ment, the glove finger is not only extruded, but, as
the extrusion is completed, the paste will be forcibly
driven from its end.

The drone is very blunt at the termination of the
abdomen, which turns somewhat under, so that the
orifice is inferior. By well-regulated finger pressure
upon the internal organs, commencing from the
thorax, as previously mentioned, the orifice becomes
more external, and, rolling out its internal wall, we
first bring into view a greyish-brown, rounded part,
the thickly-set, short spines, with curved points,
covering it, clearly indicating its purpose. This is the
“ masque” of Reaumur ( m ', E), which is simply the
side view of which m, A, is the front. The pneumo-
physes ( h') now present themselves, unroll, fill out with
air, in bubbles, as represented, and take up a position
(h') in advance of the “ masque The process con¬
tinues as the pressure is increased, the fan-shaped
appendage {/') now, like the last, turning absolutely,
so that that which was the inside becomes the outside;
and here, in nineteen cases out of twenty, the extra¬
vasation ceases. But in coition it is continued until
the bean has become external to the drone, so that
the spermatophore is lodged in the common oviduct
(co.d, Fig. 42) of the queen. Let us now endeavour

ORGANS OF THE DRONE.

205

to understand why finger pressure is usually unsuc¬
cessful in completing the remarkable process. The
theory I suggest appears to me almost conclusive.
The force which determines the change now under
consideration is derived from the pressure the drone
brings to bear upon the sexual apparatus, by a
violent contraction of the abdominal muscles. Sup¬
pose this equal to xlb. on the square inch, the con¬
tents of the ductus ejaculatorius, with every internal
part, will be subjected to the same; but the muscles
of the duct itself also contract with great energy and
power, and, if equal in force to those of the abdomen,
add another lib. pressure to the mixed mucus and
spermatozoa within. The first mechanical force tends
to drive the organ to the outside of the abdomen;
but the second drives the spermatophore backwards,
so that it is blown out like the pellet from a popgun,
and then the aperture (0) allows to pass some of the
fluid from the ductus ejaculatorius, just as air escapes
from the glove finger after the removal of the sup¬
posed mass of paste. Finger pressure fails because,
although it can fully substitute the muscular energy
of the abdomen, it cannot give any equivalent for
the driving energy of the ductus ejaculatorius; but a
little practice will overcome the difficulty, carefully
continuing to drive the body contents forward being
all that is necessary. When the last step is reached,
a sudden explosive effect is produced, which will soil
the clothing with flying droplets, unless care be taken.

The queen has no pressure within her abdomen,
because she has now no gravid ovaries, and her air
sacs are small, so that no opposition is offered to

206

BEES AND BEE-KEEPING.

the large mass of material to be transferred. What
has been said upon Flight will more fully explain
the case of the drone. The more distended the
stomach (and the male always leaves the hive, on
a love tour, loaded with honey) the more easily is
the extrusion accomplished; but it would be utterly
impossible unless the air sacs were stretched to their
utmost capacity; so that coition is impracticable on
foot. This explains why Huber never saw the accouple-
ment between drones and a virgin queen shut to¬
gether in a box, and why fertilisation in confinement
—the dream of enthusiastic apiarists—has, to this
hour, presented difficulties which would appear to be
practically unconquerable. The natural laws against
interbreeding shows this fact to be beautiful in its
fitness. The queen is not importuned in the hive,
and, when she flies abroad, the fleetest drone is more
likely to succeed in his addresses than another, and
thus he impresses upon posterity some part of his
own superior activity and energy. The slow and
weak in the race die without heirs, so that the sur¬
vival of the fittest is not an accident, but a predeter¬
mination. In previous chapters we have considered
his highly developed eyes, meeting at the vertex of
his head; his multitudinous smell hollows, and his
strong and large wings, the advantage of which now
appears in a clearer light; his quickness in discover¬
ing a mate, whose neighbourhood is to him filled with
irresistible odours, and his ability in keeping her in
view during pursuit, are no less helpful to his purpose
than fleetness on the wing ; but the success of his
suit brings the close of his career, for, quickly after

ORGANS OF THE DRONE.

207

the deliverance of the spermatophore, leaving his
abdomen surprisingly flattened and reduced, the organ
is torn from his body, in a manner respecting which
we have nothing better than hypothesis (or reputed
observation, which can hardly be regarded as either
conclusive or satisfactory). His death follows, but
certainly not so instantaneously as some have asserted,
and the queen returns to the hive, bearing at the
extremity of her abdomen the marks of her impreg¬
nation, as protruding shreds of torn membrane, to be
dragged away, dried and shrivelled, during the next
twenty-four hours. She is now more than a female;
she has within herself the potency of the two sexes,
and, during the term of her whole natural existence,
she will be able to determine and accomplish, in
time and number, within her own body, the mystical
union of male and female elements which constitutes
the act of fertilisation.

The powers of the drone just described are, almost
with certainty, not alone true for those brought up
in the normal cells of their sex, the issue of a fertilised
mother, but for all indifferently. Hereafter we shall
more fully explain that the egg yielding the drone is
unfertilised, so that those born of mothers that have
never mated (drone breeders) are as perfectly de¬
veloped and as fully virile as the others. Dwarf
drones also, raised accidentally in worker cells, or
those from the eggs of so-called fertile workers, or
workers which, although incapable of impregnation,
have yet commenced ovipositing, seem not one whit
behind the rest. Leuckart has claimed to have well
established this fact for some drones produced by an

208

BEES AND BEE-KEEPING.

Italian fertile worker, and which gave, with a black
queen, some workers of the mixed race. In such
cases as these, where so much is beyond the reach of
actual observation, it is best to cautiously abstain
from dogmatising, but the spermatozoa which these
drones contain I have found perfectly indistinguish¬
able, microscopically, from those in the normally pro¬
duced insect.

The statement that fertilisation differentiates the
sex in bees—a matter into which we shall enter fully
hereafter—introduces some of those curious freaks in
which the parts common to the two sexes are dis¬
tributed, in ludicrous confusion, to one individual, to
which the name hermaphrodite is alone applicable.
A few strange cases have occurred in my own apiary ;
but the most remarkable were sent me through the
kindness of Mr. Thompson, of Blantyre, his know¬
ledge of the typical forms enabling him to detect the
abnormal ones, which, no doubt, exist more commonly
than some suppose, but pass unnoticed. An account,
in few words, of three of these will suffice. No. i :
Head—worker; perfect worker eyes, antennae, and
tongue; Thorax—worker, except targum (back plate),
which is that of drone; legs all worker but one—
the right of the third pair ; Abdomen—completely
drone in outline ; seven segments, sexual organ male,
but actually accompanied by a rudimentary sting and
small poison bag; sting partly developed on one side,
and aborted on the other. Had died with the sexual
organs protruded. No. 2 : Head—partly worker; drone
tongue; one compound eye large, and rising nearly to
vertex, other that of worker, ocelli set far back;

ORGANS OF THE DRONE. 209

Thorax—worker, but too wide to be normal; small and
imperfect wings; very narrow plantae on hind legs,
otherwise like those of worker; Abdomen—flat and
wide, carrying imperfect drone organs. No. 3 : Head
—drone, short tongue, and eyes meeting at vertex ;
Thorax—wide, but of worker above; first pair of legs
those of drone, the rest worker; Abdomen—like that of
drone, but with only six rings. Some such bees as
these alive is a desideratum, as their internal structure
would aid in the solution of some questions of
homology and development.

Since the queen mates but once, it follows that
only an inconsiderable fraction of the drones raised
really complete the intended cycle of their being.
Colonies of bees, left entirely to their own devices,
will often produce in the spring from six to eight
thousand of these males, which consume much and
yield nothing, when perhaps but one, or at most two,
queens raised in that colony will need fertilising.
These facts, incorrectly interpreted, in the absence
of a knowledge of the beautiful laws by which these
matters are regulated, have led to the supposition
that some other office was fulfilled by the drone, he,
it has been said, being especially intended to maintain
the temperature of the stock after the swarm has
departed. It is quite fatal to this baseless theory,
that drones principally congregate on the honey, and
not on the brood cells, and that they often, in great
part, leave with the swarm. But, above all that, when
the queen has been fertilised, they are frequently
killed as useless incumbrances, no longer to be
tolerated, and the cooler the weather, as it slackens

210

BEES AND BEE-KEEPING.

the honey yield, the more certain is their de¬
struction. Mr. Haviland* has, in a very thoughtful
and well-argued paper, treated this matter. He
points out that, “If hive bees were in the habit of
producing, as most solitary bees do, males not greatly
exceeding in number the females, then the queen of
that colony which produced most drones and fewest
swarms would leave most descendants, for a queen may
leave descendants by her sons, or by any daughter who
is provided with a swarm of workers ; and it must cost
the colony far less to rear a drone than a queen, and
all the thousands of workers who must accompany
her if she is to have a chance of leaving descendants.
Hence, indeed, until the chance that a drone would
have of leaving descendants is far less than that a
queen would have, the excessive drone-producing
colonies would naturally be selected, and the selection
of variations favourable to the colony might conquer
those favourable to the species.” Paraphrasing, in
part, Mr. Haviland’s words, it is clear that the mating
queens of an apiary are more likely to meet drones
from those colonies raising them in vast numbers
than from those furnishing few. The instinct, then,
of heavy drone-production is carried into the greater
number of new colonies, an effect to be intensified
at each swarming epoch, so that there is a perpetual
tendency to increase the evil. It must, however, also
be argued, that a large production of drones is, in
one respect, favourable to the species, in that it
minimises the risk of the young queen in seeking
fertilisation ; for, the greater the certainty and prompti-
* “ The Social Instincts of Bees, their Origin and Natural Selection,” 1882.

ORGANS OF THE DRONE.

211

tude of a rencontre , the fewer excursions with the
object of mating will be necessary. The natural
check is the loss which the horde of consumers entails,
causing the colonies in which the instinct is most
highly developed to die out in times of scarcity, or
during the winter.

It is interesting to note, that the very causes which
have led to a development of drones disadvantageous
to the species, has also produced an instinct for their
destruction so soon as any chance of further normal
need of fertilisation has ceased for the season. These
pleasure-loving and lazy creatures thus come under
a general proscription when honey, or, rather, food,
is no longer yielded abundantly, for their evil day
may be put off indefinitely, by giving their stock a
constant supply ; and even sometimes when the
edict has gone forth for their destruction, a favour¬
able turn in the weather, increasing the honey yield,
will lead to their re-admission. But no sooner does
income fall below expenditure, than their nursing
sisters turn their executioners, usually by dragging
them from the hive, biting at the insertion of the
wing. The drones, strong for their especial work,
are, after all, as tender as they are defenceless, and
but little exposure and abstinence is required to
terminate their being. So thorough is the war of
extermination, that no age is spared; even drone
eggs are devoured, the larvae have their juices sucked,
and their "remains” carried out: a fate in which the
chrysalids are made to take part, the maxim for the
moment being, “ He that will not work, neither shall
he eat.”

T 2

CHAPTER XIV.

QUEEN ORGANS AND DEVELOPMENT.

Parallelisms between Queen and Drone—Both Domes¬
tically Helpless—Ovaries and Oviducts—Homologies
—Spermatheca : Microscopic Examination of —
Leidy’s and Leuckart’s Observations—Movements of
Spermatozoa—Parthenogenesis : Curious Examples
of—When the Queen Mates—Exceptional Cases —
Drone-breeding Queens—Drones have no Fathers —
Dzierzoris Experience—Fertile Workers—Inevitable
Conclusion—Appendicular Gland: Its Homologue —
Spermathecal Valve: Its Muscular System — Name¬
less Gland—Number of Spermatozoa—Old Queens
Breed Drones only — Number of Fertilisations
required — Wasps—Fertilising Pouch—Egg Struc¬
ture—Micropyle : Uses of; and Origin—Ovaries
of Workers : Barren and Fertile — Siebold’s Ex¬
aminations — Oviduct: Structure of—Prolonged
Vitality of Spermatozoa—Results of Freezing
Queens — Paralysis producing Drone-breeders —
False Theories—Worker Egg a Misnomer — Develop¬
ment of Larva—Queen Cocoon and Exuvium.

The surprising sexual development of the drone,
and his extreme domestic helplessness, are paralleled

ORGANS OF THE QUEEN.

213

by the queen (Fig. 5), which, apart from her faculty
of reproduction, is almost in every point the inferior
of the worker (see page 55). We have learnt that both
she and her partner have relatively small brain de-

Fig. 42— Ovaries of Queen, &c.

, Abdomen of Queen, Under Side (Magnified Eight times)—P, Petiole; O, O
Ovaries ; hs, Position Filled by Honey Sac; ds, Position through ’ which
Digestive System Passes ; o d, Oviduct; co.d, Common Oviduct; E, Egg-passing
Oviduct; s, Spermatheea ; i, Intestine ; pb, Poison Bag ; p.q, Poison Gland •
st, Sting; p, Palpi. B, Rudimentary Ovaries of Ordinary Worker— sp, Rudi¬
mentary Spermatheea. C, Partially Developed Ovaries of Fertile Worker—
sp, Rudimentary Spermatheea.

velopment, and that the tongue of each is so short
as to be unfit to gather honey ; that their jaws are
not suited to comb-building, and that neither has

214

BEES AND BEE-KEEPING.

wax glands; that the eyes of the queen are smaller
and less prominent than the worker’s, and her an¬
tennae inferior, both in size and organisation. Her
legs, though stronger, are less perfect, having neither
pollen baskets nor pollen brushes; while the webbed
hairs of the worker’s thorax—effective instruments
in food collection—she does not possess; her wings
are less developed, and her sting likely to be ren¬
dered useless by atrophy and inspissation of venom ;
her digestive system is less complete, and her gland
structures relatively defective, or wanting. Under
the social instinct, she, like the drone, has been
developed in one direction only; but here her facul¬
ties are more extraordinary than any to be found
outside the order Hymenoptera.

If her abdomen be cut open down the sides by fine
scissors, and the first three ventral plates and the
chyle stomach removed, we discover two very large
organs (O, O, Fig. 42), filling nearly the whole of the
inclosed space, which corresponds exactly to that
occupied by the testes in the drone. These are the
ovaries, and consist each of from 100 to 150 blind
tubes, lying side by side, and gathered into two con¬
sistent, conoid bundles, by countless small tracheae,
w T hich act as connective tissue. The ovarian tubes
are, at the upper end, very small, and here each egg
is represented by an initial cell (the germ cell), which
passes on during its development, receiving first its
vitellus, or yolk, and finally being .coated by the
chorion, or outer skin (B and C, Fig. 46). It then con¬
tinues moving downwards, as room is made for it by
the escape of the mature eggs at the lower, wider end.

ORGANS OF THE QUEEN. 215

Egg germs are far later in making their appearance
in the queen than are the spermatozoa in the drone,
the former being invisible up to the time of the
hatching of the queen nymph, whose ovarian tubes
then are filled with pellucid globules, resembling those
that precede the appearance of the seminal filaments
in the drone testis. The eggs are of a pearly-white
colour, and, during the time that queens are actively
ovipositing, more than a dozen, in various stages of
maturity, may be found in a single ovarian tube, or
follicle, standing end to end, like the beads of a
necklace. Of these, at times of activity, many will
be ready for deposition ; but, in winter, the number
in progress will be reduced to one-half, or less, while
scarcely any in a perfected condition exist. Each
tube emerges into the oviduct ( od ), the commence¬
ment of which is formed by the opened-out walls of
those on the outside of the ovary. The inner tubes
unite together at their lower edges, and so complete
the cover of the oviduct above, forming beneath them
a funnel-shaped cavity (the ventricle of the oviduct),
into which each egg first enters in its passage from
the tube in which it had been matured. The delivery
pipes of these funnel-shaped hollows (the oviducts),
uniting to form the common oviduct ( co.d ), are really
very highly organised channels, possessed of curious
powers of the greatest moment.

The similarity between the drone and queen must
here be remarked, the testes and vasa deferentia
(Fig. 41) bearing, both in structure and position, a
great resemblance to the ovaries and oviducts of the
queen, these parts being, really, respectively the

2 l6

BEES AND BEE-KEEPING.

homologues of one another, as are also the germ
cell (initial egg) of the ovary, and the sperm cell
(spermatozoon) formed in the testis. The egg, as
laid, contains not only the germ cell, and possibly
the sperm cell—the male and female elements for
the production of a new individual — but also a
store of food (food-yolk), making up its mass, and
supplying material for the development of the embryo,
until it is capable of absorbing nutrition by the
processes of ordinary digestion. We noticed, in the
last chapter, that the spermatozoa of the drone, as
developed, passed on to a store-chamber (the vesi-
cula seminalis), where these sperm cells awaited
utilisation. The homologue of the vesicula is clearly
a globular pouch in the queen (the spermatheca,
s, Fig. 42, and Fig. 43), which receives and becomes
the depository of the millions of spermatozoa ejacu¬
lated during the marital flight. Again, at the time
of mating, the spermatozoa require a medium in
Avhich they may be floated into their proper desti¬
nation, and, to supply this, the mucous gland ( mg ,
Fig. 41) is provided ; it is into this that the vesi¬
cula seminalis opens, and, during ejaculation (see
page 205), the mucous secretion and the spermatozoa
are sent forward together. The mucous gland has
also its representative, or homologue, in the queen,
in the appendicular gland (Fig. 44) of the sperma¬
theca.

To return to the queen. Near the commencement
of the common oviduct ( co.d , Fig. 41), which is fastened,
by complicated attachments, to the fifth abdominal
ring, we find the before-mentioned globular body (j),

ORGANS OF THE QUEEN.

217

rather more than ^g-in. in diameter, glistening like
burnished silver, because coated with the closest
and most densely felted plexus of tracheae with which
1 am acquainted. This spermatheca is in structural
communication with the common oviduct, but the
smallest roughness will break it from its attach¬
ment, and will frustrate any endeavour to discover
how it is filled up and used. Should it, by accident,
become detached, however, we may still study the
exceedingly curious and complicated valvular appa-

Fig. 43.—Spermatheca (Magnified Forty times).
a, Space filled by Clear Fluid; !>, Mass of Spermatozoa; c, Spermathecal Duct;
d, d, Spermatozoa in Activity.

ratus with which it is furnished. Removing it to the
stage of the dissecting microscope (see page 74), and
surrounding it with dilute glycerin, we get glimpses
of a contained membrane between the meshes of the
investing tracheae. So far as I know, those who have
studied this matter have failed to discover that these
tracheae merely closely embrace the actual sperma¬
theca, and that they in no instance enter its walls;
but such is the fact, and, by very careful teasing and
cutting with needle-knives, we may so separate the

218

BEES AND BEE-KEEPING.

multitudinous air tubes that they may be pulled off,
as a rind from an orange. The sac itself (Fig. 43) is
now seen to have beautifully transparent sides, giving
faint indications of originating in coalescing cells, but
having no discernible structure, except near its outlet,
where it has an epithelial lining. Through its sides,
if the queen is unimpregnated, we discern only a per¬
fectly clear fluid.* But should the queen have recently
mated, the whole interior is densely clouded and semi¬
opaque, since it is perfectly filled with spermatozoa,
which are recognised at once as identical with those
previously found in the drone, and from whom they
have been received and packed by a process we
can only understand hereafter; but, as older and
yet older queens are operated upon, the sperma¬
tozoa decrease in number, but, instead of being
generally diffused, are gathered into a tolerably com¬
pact mass, which lies in contact with the aperture
(c, Fig. 43), the remainder of the sac being occupied
with a transparent liquid, as in virgin queens. The
countless multitude of spermatozoa is arranged in a
definite manner, resembling a collection of long tresses
(< 5 ) combed out after recent plaiting, and as indicated in
the Figure. The extremities of the motile threads point
towards the aperture, while, from their upper surface,
spermatozoa are observed to rise in different spots
(d, d ), like microscopic eels, long and thin, curling and
twisting with much grace, as they hold on by their

* Langstroth, in notes at pages 126 and 213 of his book, tells us Leidy
found a granular fluid, and Leuckart one that was clear. Leidy is
certainly in error; while neither of these observers made, in any true
sense, a dissection of the parts, since they merely crushed the sperma-
theca flat, and examined the escaping matter.

ORGANS OF THE QUEEN. 219

tails. After a few seconds, they lapse into quietude,
to be, in turn, succeeded by others ; and, in a warm
room, this curious set of movements will be long con¬
tinued, even though several hours have been occupied
in dissecting the abdomen whence the spermatheca
have been taken.

Gently squeezing the spermatheca shows, since no
spermatozoa escape by the duct, that it is closed by
a valve, whose structure we must, by-and-by, study.
The pressure increased, the delicate bag at length
bursts, and a true microscopical marvel awaits us.
The spermatozoa escape in tufts, consisting of hundreds
of thousands, each of which is wriggling to be free;
and quickly they are widely spread, curling and un¬
curling with a peculiar snapping movement, and with
an energy that baffles description. Their powers in a
few minutes begin to wane ; then, one after the other,
they take a form closely resembling two 8's, one over
the other, surrounded by a rather larger O ( a , C, Fig.
41). When all have sunk to rest, this singular pattern,
repeated with strange regularity, covers the field,
though sometimes the threads take a wider outline,
as the illustration makes clear. It remains to be seen
by what means these spermatozoa are packed in the
spermatheca after being received from the drone, and
how they are transferred to the egg as required, and
whether they are so transferred in all cases. But
before considering the structures involved, it is well
that we should direct our attention to the theory of
parthenogenesis, or production by a virgin, which facts,
observed half a century since, satisfactorily showed to
exist both in wasps and bees; but the argument

220

BEES AND BEE-KEEPING.

remained entirely constructive in character, until I was
fortunate enough to establish for it an anatomical
basis,* which not only explains the facts, but the
structures which make them possible.

Parthenogenesis, or reproduction wuthout fecunda¬
tion, by virgin and perfect females provided with
ovaries and spermatheca, is no new fact in entomology.
It received recognition at first in the earlier half of
the eighteenth century, in the case of some virgin
female silk moths, and afterwards in others of the
Bombycidaa , all of which produced eggs hatching out
both sexes. Later, an incomplete parthenogenesis was
observed in the Psychidae and some nearer relatives
of the bee (the gall flies), the virgin females laying
eggs yielding exclusively females (the less perfect
form in these genera) ; the process being repeated,
during twenty generations, without a single male in¬
dividual presenting itself, or one case of impregna¬
tion having occurred. Indeed, amongst some moths,
the male is at present altogether unknown. Nor have
we at all exhausted our knowledge of these surprising
variations from a rule formerly thought to permit of
no exception; for, amongst other similar cases, in a
species of Cecidomyia , a small insect, living, in the
larval state, beneath the bark of trees, the larva is
itself fertile, producing creatures in its own likeness,
which at maturity tear open the side of the parent
and escape, themselves to similarly give rise to another

* “ The Apparatus for Differentiating the Sexes in Bees and Wasps.
An Anatomical Investigation into the Structure of the Receptaculum
Seminis and Adjacent Parts.” F. R. Cheshire. Journal Royal Micro¬
scopical Society, February, 1885.

ORGANS OF THE QUEEN.

221

brood, until the close of the season, when true meta¬
morphosis occurs, and the adult form of the insect
makes its appearance.

And who, too, is unacquainted with the far too
common, sexless, budding Aphis , passing through
several generations, until perfect, sexual, wingless
Aphides are brought into existence, upon which seems
to be laid the task of continuing the race to the
succeeding year. And, leaving the domain of insects,
we meet with no less curious instances. Amongst
the lowly Rotifers, by example, females are generated
by virgins, and males by mated individuals. Nor is
parthenogenesis unknown to the world of plants. Dr.
Asa Gray gives, as an example, Caelebogyne, respect¬
ing which he says: “ Parthenogenesis is thus con¬
firmed, and is known to occur in most polyembryony.”*

But it may be argued, that the queen bee is only
capable of filling her office as a mated insect, and
that, consequently, these illustrations do not apply.
In the majority of cases, this is so. Ordinarily, for
the first six days following her escape from the
queen cell, she manifests no disposition to make an
excursion abroad, although numerous drones may
be without, floating in the bright sunshine; and
even after this period, when the elements are un¬
favourable, through chilly winds or falling rain, or
in the morning or evening, when drones are at
home, she quietly stays within ; but, at the age
named above, during the three hours or so which
follow midday, when the weather does not forbid,
and when the drones are executing their sonorous
* Structural Botany,” page 285. Macmillan and Co., 1880.

222

BEES AND BEE-KEEPING.

evolutions, the young queen, if prevented from
leaving, becomes greatly agitated, seeking an exit at
every point, until the drones are once more at home.
If at liberty, she flies daily with increased anxiety,
until the object explained in the last chapter has
been realised, when, about forty-eight hours after, she
deposits her first eggs, which invariably produce
workers.

But exceptional cases often arise, and it is in har¬
mony with facts such as those before given, and
which have long been known, that a queen, or
mother-bee, is not doomed to total sterility if raised
at a part of the year when drones do not exist, but
that she, although later than at the normal period,
begins to deposit eggs, which, however, are in no
instance converted into workers, but invariably pro¬
duce drones, which must, of course, in her case at
least, be generated parthenogenetically. Queens
having defective wings, and so incapable of mating,
are also invariably drone-breeders. Similarly, if a
queen of the Italian race {Apis Ligustica ), which has
consorted with an Italian drone, be placed in a hive
containing English bees (Apis Mellifica) only, and
which is itself located in a neighbourhood where
Italians are unknown, all her progeny, both workers
and drones, will, to the end of her life, continue
pure, carrying their characteristic yellow abdominal
bands, and a thousand other minor distinctive pecu¬
liarities; but should she leave with a swarm, or
die, the workers will raise a successor from one
of her eggs. The new queen of unmixed blood
must of necessity mate with an English drone

ORGANS OF THE QUEEN.

223

(allowing, for the sake of the argument, that
her mother has produced none), and, as a con¬
sequence, the workers, her progeny, will partake of
the qualities of the two races, exhibiting among
themselves those variations for which hybrids are
remarkable. But her drones, on the contrary, will still
be absolutely Italian, again showing that, although
their mother was impregnated, hep impregnation had
in no way influenced their generation, or that they,
as before, had a mother but no father; so that the
eggs whence they came had in some way escaped
fertilisation. Almost all apiculturists have had abun¬
dant evidence of a kindred kind; but it was the
introduction of Apis Ligustica into Silesia, in 1853,
which gave Dzierzon the first incontestable proof of
the parthenogenetic production of the drone. Yet
further evidence is given by the occasional appear¬
ance of fertile workers, whose existence has . been
previously referred to, which, from their anatomical
structure, are incapable of coition. These, never¬
theless, deposit eggs which, for reasons now evident,
produce drones only. The conclusion cannot be
evaded, that, in the genus Apis, where the least-
developed form would appear to be the drone, the
egg is already sufficiently vitalised for giving him
birth when it has reached maturity in the ovary,
but it requires the concurrence of the male sperma¬
tozoa to produce the female, the most highly endowed
and organised of the sexes amongst the Hymenoptera .
Our normally mated queen, then, according to season
and the necessity of the colony, deposits eggs, either
in the smaller cells, yielding workers, or in the

224

BEES AND BEE-KEEPING.

larger, furnishing drones, because she possesses the
extraordinary faculty of giving at will, or withholding,
spermatozoa from the egg about to be deposited.
Let us return to the study of her anatomy, for the
purpose of unravelling the mystery of the mechanism
by which this is accomplished.

sp, Spermatheca deprived of Tracheal Coat; c, duct; a, a, a, b, b, b, Bight and
Left Branches of Appendicular Gland; t, t, Duct of Same ; d, d, d. Nerve
Connections; e and /, Sphincter Muscles; q, Muscle to Extend Valvular
Opening; h, Muscle for Closing Valvular Opening; i, Ganglion Lying Under
Muscle; 7c, 7c, Duct for Spermatozoa; l, l, Glandular Structure surrounding
Duct.

Taking a complete spermatheca, we turn it until
it presents an outline not unlike the back of a
man’s head, carrying a pair of large and prominent
ears. The latter are the upper ends of the right
and left branches of a gland ( a , a , a , b, b, b, Fig. 44,
and ag , Fig. 45), which are of considerable length,
and about y^-in- in diameter, and which are held in

ORGANS OF THE QUEEN.

225

position by receiving very numerous twigs from
the tracheal net (sp } Fig. 45) inclosing the sperma-
theca. These branches pass down the opposite sides
of the sac, and unite near to its aperture ( c , Fig.
44). The whole gland consists of nucleated cells,
surrounding a common duct (/, /), which runs from
end to end, and enlarges somewhat during its course.
Its type is very distinctly intracellular (page 77),
and the different ductlets, many thousands in number,
leading from the very numerous independent cells, are
indicated in the Figure. Its activity and importance
are further sh,own by the multitudinous nerve twigs
and cells ( d , d , d), giving it general energy, and
bringing its various parts into relation. The name
“appendicular” appears to me ill-chosen, since there is
every reason to regard this gland as the homologue
of the mucous gland of the drone; “ mucous gland
of the female ” would have been, therefore, more ex¬
pressive. New names are often confusing, so the old
one will be retained during the description. The
spermathecal duct ( c ), which is short, stiff, and
slightly ribbed, points towards, but does not im¬
mediately enter, the duct of the appendicular gland.
I find the disposition of the whole to be that of a
valve, to which, and to these ducts, are attached five
main muscles, two being sphincters (indicated at e and
f). The latter extend upwards farther than repre¬
sented, their continuation being omitted lest they should
obscure the structure of the valve before and behind
which they actually pass. These sphincters are the
instruments for respectively and independently closing
the appendicular-gland and spermathecal ducts. They

226

BEES AND BEE-KEEPING.

are separated by an intervening wedge-shaped disc,
so that they lie towards each other, at an angle
of from 30° to 6o°, and may be beautifully shown
by polarised light. An indurated integument, pro¬
bably a chitinous plate («), is pushed towards the
spermathecal duct, by the contraction of its proper
sphincter ( e ), and in this work it is aided by the
muscle h, which is one of two, whose tendinous
extension is about 10 1 00 in. in diameter, or -^th the
thickness of a human hair. These muscles would, no
doubt, all remain tense while the insect was in a con¬
dition of repose ; but should she be engaged in
ovipositing, and spermatozoa be required for fertilisa¬
tion, the muscle g, by contraction, would lift the plate
lying above and between 0 and k , to which, by a
complex tendon, it is attached. Into the cavity (1?)
thus opened, spermatozoa would pass; the two
sphincters at the same moment relaxing, an outflow
of glandular secretion, as indicated by the arrow,
would be ready to sweep' the spermatozoa towards
their destination in the common oviduct, and all
would be driven on by the appendicular sphincter
e first contracting, followed in order by the second
sphincter (f), and muscle marked h, when both
ducts would be closed, and the repose condition
re-established.

A most remarkable adaptation here arises. The
spermatozoa yielded by the drone are, probably, not
usually more than 4,000,000 in number. It is, of
course, extremely difficult to make a calculation ;
the very highest estimate I have ever reached is
12,000,000 ; Leuckart states that the spermatheca may

ORGANS OF THE QUEEN.

227

contain 25,000,000 of spermatozoa. While not denying
possibility to his estimate, I certainly think it far
too high. Whichever sum be accepted as the true
one, it is demonstrable, that economy in the distri¬
bution of these fertilising threads is of the highest
possible moment, for, should they be shot forth hap¬
hazard, they would be exhausted long before the
queen’s death, when she would be, of course, reduced
to the condition of a queen that had never mated,
and so become, like such, a drone-breeder: a circum¬
stance by no means uncommon—presenting itself,
indeed, quite frequently where, under careless manage¬
ment, queens are allowed to fade out instead of being
displaced. They may then, in the absence of accident,
attain the ripe old age of four, or even five, years.
Many of these ancient dames — discarded because
they no longer yielded workers, or only a few, amidst
many drones, and these produced in worker cells—
have been sent to me for dissection, and I have inva¬
riably found the spermatheca quite denuded of its
spermatozoa, or only containing such a miserable
residue as to clearly show that the eggs could, at
the best, be but occasionally fertilised. The economy
we see to be so essential is secured as follows: The
duct ( k, k) through which the spermatozoa pass, as
extruded in detachments from the spermatheca, I find
to be the centre of another gland (/, /), which seems
to have escaped the attention of previous observers.
This gland we may fairly infer to be excited to
secretion by the presence of the spermatozoa, just as
food excites our salivary glands to the secretion of
saliva, and the stomach to the secretion of gastric

u 2

228

BEES AND BEE-KEEPING.

juice. Spermatozoa, thickly present, will cause the
addition of large quantities of fluid, more widely
separating them. Their absence (for this gland is
most richly provided with nerve twigs, which send
numerous loops to the muscles previously described,
and to the ganglion, •/, lying under the muscle, g,
and placed just over k in the Figure) will yield the
action which will send a new contingent forward as
I have described, and so they come to be paid out
with some regularity. The necessity for this regu¬
larity will be better appreciated if we remember that
a prolific queen will lay, during her life, 1,500,000*
eggs (see page 83)—a number so vast that the eggs,
lying in contact, end to end, would stretch about
one and three-quarter miles. Deducting a few thou¬
sand for drones (for the production of which sper¬
matozoa are not needed), the remainder would each
require an independent fertilisation, and, for this work,
possibly, 4,000,000 spermatozoa, or even less, may be
at command. In this connection, it is most interest¬
ing to note that the spermatozoa, in the different
genera and species, stand in beautiful relation to the
number of eggs deposited by the fertile female. In
the queen wasp, by example, the fecundity is much
less, happily, than in the honey bee, and so the sper-
matheca is considerably smaller, the capacity of that
of the former insect being only about one-fortieth of
that of the latter, the spermatozoa being nearly of
the same size. The organs of the male wasp are
correspondingly reduced.

# This number is much beyond an average; but it certainly has been
reached, if not exceeded.

ORGANS OF THE QUEEN.

229

I have found the channel k, k, to contain a mem¬
brane of extreme tenuity, only made visible with
difficulty, and this is remarkably convoluted, after the
manner of the epididymis of higher animals. Tracing
the channel onwards till it perforates the side of the
common oviduct turned from us in Fig. 42, or towards
us in Fig. 45, a bifurcation is detected, with one
channel, apparently wide and indefinite, which is

Fjg. 45.— Passage Connecting Oviducts with the Exterior of the Body
(Magnified Ten times).

sp, Spermatheca ; ag, Appendicular Gland; a, Upper, and d, Lower Path for Eggs ;
be, Bursa Copulatrix ; p, Fertilising Pouch; m, m', Muscles for Contracting
Side Path.

quickly lost by its becoming confluent with the lower
part of the oviduct, whilst the other enters a
central and curiously-folded apparatus ( p, Fig. 45),
which, for a reason to be presently given, I shall
denominate the “ fertilising pouch.” I have strong
reasons for supposing that the path from the bursa
copulatrix (be )—into which the male organs are
locked by the horned pneumophyses (see page 203)—•

230

BEES AND BEE-KEEPING.

and from the parts of the oviduct above it, through
the deeply-folded pouch (/) to the spermatheca, is so
involved, that it would not be possible for the sper¬
matozoa, by following it, to enter the latter when
given up by the drone; but that, in the early life
of the queen, the second wider and straighter
channel to which I have referred, is fully open,
and by it the spermatozoa, with their inscrutable
power of self-direction, pass upwards, avoiding the
mazes of the fertilising pouch, and packing them¬
selves for future use pretty much as they were
arranged in the spermatophore in the drone’s body.
The queen, if still unmated at four or five weeks old,
becomes incapable of copulation, or, at least, she
evinces no desire for it, which fact possibly marks the
time when this lower passage closes, such closure, in
a mated queen, forcing the spermatozoa, in descend¬
ing , to take their way by the fertilising pouch.

If a central comb be lifted from a hive during
the summer months, eggs in number will be dis¬
covered. If one of these be removed from either a
worker or a drone cell, by means of the wetted point
of a camel-hair pencil (for they are deposited with a
secretion covering them, which causes them to adhere
by the end, as at A, Fig. 46), its surface will be
found, if examined microscopically, to be covered by
a beautifully reticulated membrane (the chorion, B and
C), almost as though a tiny pearl had been covered
with what the ladies call blonde, many hundreds of
the meshes of which are required to coat it completely.
Arranging the egg so that we get a view of the larger
end (D)—for which nothing excels a -|-in. objective

ORGANS OF THE QUEEN.

231

and Lieberkiihn—we find the netting disposed in a
radiating pattern, reminding one of the cordage over
a balloon, which leads up to the strong ring at top.
In the centre lies a single aperture (the micropyle)
marking the point for the insertion of the funiculus,
by which the egg was attached during its growth,
and from which it separated itself when sufficiently
matured. The minuteness of the opening does not
prevent its being continued through the underlying

Fig. 46.—Bee-Egg and Details.

A, Position of Eggs at Base of Cells (Natural Size). B, Egg (Magnified Twenty-five
times), showing Reticulated Chorion—c, Base attached to Cell Bottom;
ma, Micropylar Aperture. C, Chorion (Magnified 200 times). D, Micropylar
Aperture (Magnified 100 times).

egg membranes, and giving an opportunity of entrance
to the spermatozoon, whose rhythmic movements, as
though guided by intelligence, conduct it to the
micropyle when the egg passes within the fertilising
pouch, on its road towards being laid in a worker
cell. The wondrous thread enters, coalesces with
the germ, brings about fertilisation, and effects the
resulting sex, as previously recited facts force us
to believe. The egg so impregnated yields a female,

232 BEES AND BEE-KEEPING.

which will possess qualities both of father and
mother; so that the tiny spermatozoon not only
differentiates the entire creature, but communicates,
unerringly, differences of species, or even mere
variety. The spermatozoa from Cyprian, Carniolian,
Italian, and English bees are to the most refined
microscopical examination identical, and yet they
contain differences which determine almost countless
variations in form, colour, size, instinct, capability,
and temper. In 1884 I made the extremely in¬
teresting discovery, that spermatozoa, when within the
spermatheca, are subject to disease (see Diseases),
and, in one instance, in which hermaphrodite (page
208) bees occurred, this disease obtained in the
queen. Examples being so sparse, and the difficulties
of examination so great, it is not likely that this
fact will lead up to any generalisation ; but it is
most tempting to a spirit of speculation. If a sper¬
matozoon converts that which would, in its absence,
have been a male, into a female, may not a defective
spermatozoon only in part produce the change, so
that a mixed gender results ? So far as we know,
it is certainly in agreement with the evidence to
admit the possibility.

That the spermatozoon enters the egg is certain,
for it may be found, if the latter be carefully ex¬
amined immediately after deposition. Siebold,* by
crushing eggs which had immediately before been
deposited in worker cells, was the first to discover

* Siebold “On True Parthenogenesis,” p. 85 et seq., and “ Parth4no-
genese chez les Insectes” (Annales des Sciences Naturelles, 4 me S6rie,

ORGANS OF THE QUEEN.

233

the spermatozoa within. In some instances, he
thought he saw as many as three that had passed
the micropyle; but I cannot forbear expressing
the opinion that possibly, or, rather, probably,
Siebold has been in error here, since there is
good reason for imagining that one completes the
process of fertilisation. Positiveness would be much
out of place; the whole investigation is so extremely
exacting, and needs, for its successful prosecution, the
concurrence of so many favourable conditions, that
errors can hardly be avoided; the remarkable length
of the body of the spermatozoon—about y^-th of
an inch, which is more than 300 times its greatest
width—necessitates many convolutions, and would
make misconception easy. Whether we have seen
only one or more, may, for the moment, rest; but
the interesting point lies in this, that the most
careful examinations made by Siebold, and which I
have confirmed by prolonged observations, show that
no trace of a spermatozoon is found either within or
upon the eggs laid by a fertile mother in drone cells.

Dr. Donhoff claims a curious corroboration by
artificially impregnating, in 1855, an egg laid in a
drone cell, by placing upon it a little diluted fluid
from a drone testis, and transferring it to a worker
cell. Others have failed in this experiment, but the
argument for the parthenogenetic production of
drones can well afford to do without the evidence
it would supply, even if repeated by many observers.

The head ( h , C, Fig. 41) of the fertilising filament
is very narrow, that the micropylar aperture may
be passed, but, to effect this, time must be occupied ;

234

BEES AND BEE-KEEPING.

and how is this given ? My previous explanations
have made evident that the spermatozoa glide, not
into a plain tubular cavity to meet the descending
egg, but into a pouch contrived of curiously formed
folds of the lining membrane of the common
oviduct, and which, if stained with picro-carmine,
takes up picric acid and becomes yellower than the
oviduct proper, whilst its surface is dotted over with
linear patches of setae (or bristles), from two to six
in a patch, and from - 10 ^ 00 in. to g^ in. in length.
Its structure is particularly difficult to examine, but
it has three main cross duplicatures (p, Fig. 45) of
an extremely attenuated membrane, which give to
it somewhat the form of three joints of a lobster’s
tail, while it is only slightly wider than the diameter
of the egg; and I have little doubt that here the
latter is delayed when a female is to be produced,
and brought into contact with spermatozoa delivered
into the right position by the channel k , k (Fig.
44), whilst the eggs from which drones are evolved
are carried down the path ( d , Fig. 45) by the side
of the pouch to the termination of the duct, and so
escape all contact with the fertilising threads.

The oviducts are highly organised, containing a
most beautiful system of longitudinal and transverse
muscular fibres, repletely provided with nerve
twigs, evidently giving to the oviducts the most com¬
plete control of the eggs which are to pass through
them, while they are not without strong indications
of two specialised paths (b and c ), one towards the
fertilising pouch and the other to its side. Near the
junction of the oviducts, also, there are two thin

ORGANS OF THE QUEEN.

235

muscles ( m, m !, Fig. 45), for which I can conceive of
no purpose, unless it be to so reduce, by their con¬
traction, the opening lying by the side of the fer¬
tilising pouch ( p ) that an egg could not, except they
are relaxed, pass in this direction, and so escape
fertilisation. That these parts have great regulating
capability, and are not mere tubular conduits, is
proved as much by their nerves as by their muscles.
The last abdominal ganglion lies immediately be¬
neath, and in contact with, the oviducts and sperma-
theca, and, from it, branches of nerves run in
abundance into the oviducts, the spermathecal valve
muscles, the sting, and their palpi; while small
ganglia are distributed in profusion, a considerable
one lying over the valve, and sending branches for¬
ward into the fertilising pouch. The manner in which
the spermatozoon itself finds its way is utterly in¬
scrutable. The fact of its continued vitality with no
distinguishable change, either in size and form, or
motile activity, during the whole of the queen’s life,
save from five to ten days, between which ages she
usually mates, is most surprising. Constant nutrition
and oxidation can alone be capable of sustaining it
to the last in the freshness it had when first intro¬
duced to the spermatheca. Cold, however, kills it.
Here Dzierzon’s experiments have the deepest in¬
terest. He found that a queen which had been
refrigerated for some time, although capable of re¬
vivification by warmth, never afterwards laid other
than drone eggs, whilst before she had been a good
producer of workers. Berlepsch placed three queens
for thirty-six hours in an ice-house; two died, but

236

BEES AND BEE-KEEPING.

the third recovered, and laid abundantly, but drones
only resulted. Similar experiments are also related
by Langstroth, who adds that a short exposure to the
intense cold of a mixture of ice and salt will answer
every purpose. But, while the spermatozoa retain
their energies—unless means be taken to destroy
them—the queen, even when young, may become
incapable of distributing them ; and I have had, under
the dissecting-knife, not less than four examples,
that had been sent me as drone-breeders, but which
I found to contain an abundant virile supply.

This condition may arise from the spermatozoa
choking the duct, or from failure of the last ganglion,
and Dr. Donhoff relates that he has produced it
by simply gently pinching the terminal abdominal
segment with a pair of forceps, after which the
queen exhibited difficulty in laying, in consequence
of the nerve injury.

The fecundation of the mother-bee was the sub¬
ject of many false hypotheses before the facts were
discovered. Swammerdam, observing a strong odour
from the drone, supposed that this, permeating the
body of the female, fertilised the eggs. The number
of drones seemed to be explained by this aura
seminalis theory, since a crowd would be required to
produce the supposed emanation in intensity. Huber
completely overturned this fancy, by placing the
drones in a box pierced with holes, the vapours
from which left the queen a drone-breeder. De
Braw, observing little white masses at the bottom
of the cells (really the last cast skin'of the matured
chrysalis), announced that drones fecundated the eggs

ORGANS OF THE QUEEN.

237

after they were laid, after the fashion of fishes.
Huber, assured of the falsity of the idea, shut up
all the drones of a hive, letting the mother fly, and
found that she became fertile. We may smile at
these ancient blunders, but really mistakes as grave
and less excusable now obtain. It is even yet
asserted by some, as the echo of a bold guess made
long ago, by an American apiarist of just repute,
though but little acquainted with scientific matters,
that the narrower cells in which worker bees are
raised, by pressing upon the abdomen of the queen,
were the effective agents for forcing out the sper¬
matozoa, and so causing the eggs to be fertilised.
This notion, so repellent from its bald crudity, is
shown to be utterly without foundation. Not only
do queens lay worker eggs in cells whose sides
are only commenced, so that pressure cannot be
exerted, but experiment proves that the sex is
determined according to the needs of the colony.
Although the queen, if left in undisturbed possession
of the combs the workers have built, will select the
cells, and lay eggs appropriate to their sizes, she
will, if provided with one kind only, deposit in part
eggs of a sex opposite to that for which the cells
are suitable; for, if a hive be filled with drone comb,
workers will be raised in it. Fertilisation, now that
we have so far conquered its modus operandi , is
seen to be absolutely under control, and the out¬
come of a beautiful and marvellous mechanism.

But the egg, having been fertilised, may, accord¬
ing to subsequent treatment, yield either worker or
queen. To be the latter is rarely its destiny, and so

2 3 8

BEES AND BEE-KEEPING.

it is commonly called a,, “ worker egg,” which is
clearly inaccurate, and comes, like other mistakes of
this kind, from terms being introduced and made
current before the objects named are scientifically
understood. The two essential forms of egg are the
impregnated and unimpregnated, yielding the neces¬
sary concomitants of reproduction, the female and
the male, the queen and the drone; and from the
former, as the social instinct has been developed, the

Fig. 47.—Larva and Chrysalis of Hive Bee (Magnified Four times).

A, Larva (full grown)— os, b, c, d, Terminal Segments ; 1, 2, 3, 4, 5, Segments below
Head; l, Budding Legs; w, Ditto Wings. B, Condition of Change inter¬
mediate between Larva and Chrysalis; Lettering as before. C, Chrysalis ;
Lettering as before.

worker has been produced. Labour has been divided.
The queen has lost her domestic arts, which the
worker possesses in a perfection never attained by
the ancestral types ; while the worker has lost her
maternal functions, although she still possesses the
needed organs in a rudimentary state. Ovaries she
has, but so tiny as only to be found by elaborate
dissection. They escaped altogether the vigilance
and skill of both Swammerdam and Reaumur, and

ORGANS OF THE QUEEN. 239

are little better than an attenuated string of tubes,
ten or twelve in number, and destitute of eggs or
germs (B, Fig. 42), where, even yet, an indication
of spermatheca ( sp ) remains. Although the cavity of
the latter is almost entirely obliterated, the vestiges of
the appendicular gland pass into its base after the
manner of arrangement in the fertile mother. The
vagina lies at the side of the intestinal opening, and
is frequently imperforate, while the bursa copulatrix
(be, Fig. 45) does not exist, so that the reception of
the male organ is impossible. Workers, like queens,
pass through a very considerable range of variation,
and an instance of worker copulation, which has
been scientifically verified, is on record. Under ex¬
citement, and in the absence of a queen, the ovarian
tubes will, in rare instances, extend, and eggs be
laid, producing, as we now know, drones, the
ovaries, when dissected out, presenting then the
appearance of C, Fig. 42.

In previous chapters, we have traced the develop¬
ment of the larva within the egg, and have studied
the remarkable transformations in the arrangement of
the internal parts that occur during the chrysalis stage ;
and it seems fitting that we should now direct our
attention to those previously omitted transformations
which gradually change the egg, whose qualities we
have just investigated, into the outline of queen,
worker, or drone. The oviducts are provided with
secretion cells, which coat the egg with an agglutina¬
tive body, so that, as it leaves the queen, it adheres
by its smaller end, as before pointed out. It sustains
its outstanding position for the first day, but then

240

BEES AND BEE-KEEPING.

gradually sinks. The chorion of the egg (C, Fig. 46)
breaks, usually after three days (the time varies
according to temperature), and a footless larva, with
thirteen segments, exclusive of the head, alternately
straightens and bends its body to free itself of the
envelope. It is extremely curious that, before hatch¬
ing, the larva presents rudimentary legs, which dis¬
appear—a fact which some have supposed to indicate
(atavism) a reference to an ancestral type in which
the larva bore feet; but this does not seem to be
valid, for reasons which would encroach too much on
our space Towards the end of the larval period, the
three segments following the head (1, 2, 3, A and
B, Fig. 47) have little scales (/) beneath the skin
on the ventral side, which are the beginnings of
the legs, and which cannot be seen until the creature
has been immersed in alcohol; the budding wings
{w) outside these, on segments 2 and 3, are, by the
same treatment, brought under view, as are also the
rudiments of the sting in queen or worker larvae
(Chapter XII.), the male organs appearing in that of
the drone. After sealing, the fourth segment begins
to contract, and the fifth becomes partly atrophied,
so that, soon, the former constitutes only a partial
cover for the base of the developing thorax, and the
petiole between it and the abdomen, while the
latter becomes the narrow, first abdominal segment.
At page 196, it has been explained that the last
three segments disappear in forming the sting; and
now we find the fourth forming the petiole, leaving
nine of the thirteen original segments, of which three
go to the thorax, and six to the abdomen.

QUEEN DEVELOPMENT. 241

A point in relation to the behaviour of queen
grubs, as differing from that of others, chiefly referring
to the facility with which the cell containing the
queen nymph can be torn open, here requires careful
attention, because it is so generally mis-stated. After
the spinning of the cocoon, which in no case extends
far down the sides of the cell, the worker or drone
larva, as before mentioned, turns and throws up the
bowel lining and contents, and casts its skin, which,
by the creature’s movements within the cell, becomes
plastered to the walls, and joins the cocoon near
the mouth end. The legs, wings, and advancing
male organ or sting—depending on the gender of the
grub, and which before could not be seen without
treatment—are now fully visible, the name chrysalis,
or nymph, being properly applied; the modelling
continues, dimplings are seen, rounded forms become
angular, the external skeleton gathers in density and
colour, bristles appear; every organ is advancing,
and, ere long, the imprisonment and the darkness
are left for the heavenward flight of a new life,
which gave to the ancients the name and the type
of a resurrection. Huber,* especially with regard to
the structure of the cocoon, fell into errors, from
which he drew false deductions that are being still
repeated. I give a free translation, in order to
condense his meaning, which runs thus: “The worker
and drone grubs form complete cocoons— i.e., the latter
are closed at both ends, enveloping all the body. The
royal grubs, on the contrary, make cocoons which
are imperfect, being open at the posterior part,

“Nouvelles Observations sur les Abeilles,” page 218, vol. i.

242

DEES AND BEE-KEEPING.

enveloping only the head, the thorax, and the first
ring of the abdomen. Th’s discovery has given
me/' continues Huber, “ extreme pleasure, because it
evidently shows the admirable way in which Nature
has brought into agreement the different actions of
bees. Queens have a great mutual aversion, blood¬
thirstily seeking one another’s destruction. When
there are several royal nymphs in a hive, the first
one to hatch throws herself upon the others, and
pierces them with thrusts from her sting. But she
could not succeed if the nymphs were inclosed in
a complete cocoon, because the silk is strong, and
the cocoon of a close texture, which the sting would
not penetrate; or, if it penetrated, could not be
retracted, so that the queen would die the victim of
her own fury. In order that a queen may succeed
in killing her rivals in their cells, it is needful that
their hinder part be uncovered, for it is only here
that the dart will penetrate them, the head and the
thorax being clothed with strong, scaly plates. The
royal grubs should, therefore, furnish incomplete
cocoons.”

Is it true, then, Mons. Huber, that an unpoetical
little grub emulates Cassius, when, in a supreme
moment, he exclaims: “ There is my dagger, and
here my naked breast”? Iteration and reiteration,
by author after author, since your admirable investi¬
gations notwithstanding, it is utterly incorrect. We
should have had fewer writers, or more investigators
for the microscope, since the introduction of the
achromatic objective, between fifty and sixty years
ago, has been fully equal to showing that no cocoon,

QUEEN DEVELOPMENT.

243

as already said, extends much beyond the cell mouth,
the remainder of the covering of drone and worker
being the cast skin; but, in the case of the queen
grub, whose cocoon is really more extensive, and
decidedly tougher, than that of the other inhabitants
of the hive, the royal jelly, occupying the upper part
of the cell {a, Fig. 48) clearly prevents the usual method
of proceeding. The skin and bowel are, indeed, cast
as by the worker, but they are not spread out on the

Fig. 48.— Comb and Queen Cells (Magnified Twice).
a, Queen Cell, Cut to Expose Royal Jelly and Grub at Upper End; b, Thickness
of Cell; c, Dimpling Outside Cell; d, Spot where Bowel Contents and
Exuvium are Placed

cell wall. The bowel, relatively small, and containing
little waste product, is thrown against the side of the
cell at d, Fig. 48, just below the mass of royal jelly ;
and here the skin of the body is placed also, where
both can always be found, by opening a queen cell
on the third day after sealing. During the earlier
part of the changes, the developing insect adheres,
by the dorsum, to the wet royal jelly, and probably

244

BEES AND BEE-KEEPING.

continues to take nourishment through a part of the
skin. We thus see that, when the murderess arrives,
nothing but the thick cell side, of impure wax, inter¬
venes between her and her victim; the cast skin,
which is extremely delicate, is absent as a lining, it
is true, but the cocoon is placed as in all other cells.

Huber investigated the periods of evolution for the
sexes—matters, the details of which must be treated in
our Practical Section ; but his results, which are con¬
stantly given in bee books, are only approximations,
for, directly we attempt to systematically follow up the
inquiry, we find that considerable variations present
themselves : in one hive the worker bees gnawing
out, on an average, on the twentieth day after the
egg is laid, and in another not until the twenty-first;
eggs in the same hive, and of the same hatch,
especially those of drones, taking unequal times for
evolution : while the seasons of the year also make
a difference. Huber found that queens required six¬
teen days to mature, workers twenty-one, and drones
twenty-four. But even within the hive, I find the
queens can, by management, be delayed so as to
require nearly eighteen days, the workers twenty-
five, and the drones twenty-eight; while, by re¬
moval from the hive, a more considerable retardation
may be occasioned. The more rapid normal de¬
velopment of the queen is highly interesting, and
its reason is evident. As queens fight, the first
hatched has the best chance of being the survivor,
so that there is a constant selection in favour of
those rapidly maturing. There is, in a modified
degree, a similar selection of workers, for any in-

QUEEN DEVELOPMENT.

245

crease in the time occupied before they leave the
cell would heavily handicap the stock, and so
decrease its chance of sending off a swarm, thus pre¬
venting the mother from leaving descendants by her
daughters; and, since bees do not generally produce
drones until swarming has for themselves reached
probability, she would also have less chance of leav¬
ing descendants by her sons.

In closing this chapter, which also terminates one
section of our studies, we must be impressed with
the mysterious division of labour between queen
and worker, the latter fitted to honey and pollen¬
gathering, wax-secreting, comb-building, nursing,
and cleaning, with every tool she can need; the
former, for the duties of maternity, and for these
alone, with generative organs fully equipped for the
enormous work demanded of them, but that at the
expense of all those parts which minister nothing
to her proper functions ; all originating, too, in
the coalescing germ and sperm cells, endued, so
far as the eye of the body or of the mind can
carry us, with powers to build up, differentiate, and
arrange a mechanism which, though tiny, can make
us all exclaim, “ We are but of yesterday, and know
nothing!” And let us not here fall into a mistake
far too common. No natural object receives attention
that does not grow in wonderfulness under the opera¬
tion, and so each one is prone to think that his par¬
ticular subject of investigation has more in it than
any other. We naturalists and bee-keepers, as we
watch and study the little insect that has given so
much delight, are in no little danger of coming to

246

BEES AND BEE-KEEPING.

believe that we are contemplating the very master¬
piece of creation, and that we have before us a con¬
centration of wisdom and of wonder for which we
should look elsewhere in vain. It has, by example,
almost become a fashion to tell us, as a modern
manual does, in reference to the production of queens,
that “ we have here a fact which has no parallel
in natural history.” A broader view will show, as
we have just seen, that not only is this untrue, but
that quite as surprising and unlookcd for methods
of sexual differentiation not infrequently occur. Our
very mistakes may help us upward ; for should not
the fact that the things which at first we think little
we afterwards discover to be great, and that the
more we study, the more we find there is to learn,
rather prove to us the unwisdom of supposing we
have already unfolded the greatest of all wonders,
teaching us that, as yet, we only discern few marvels
where there are many, and that, did we know Nature
as she is, we should see neither less nor greater, but
fulness of beauty everywhere, the exponent of a wisdom
past finding out? The oppressive infinitudes of as¬
tronomy, and the equally inconceivable minuteness
revealed by the microscope, are but two phases of
the frame of the universe, which has touched infinity
at every point. Already, indeed, we get glimmer¬
ings that the recognition of this fact will be a goal
of science, which is now opening up to us, that not
only animals and plants have their wonders, but that
the very atoms are miracles of form and force, bound
together by relationships which are endless.

CHAPTER XV.

BEES AND FLOWERS MUTUALLY COMPLEMENTARY.

Outline of Subject—Old Errors and Modern Dis¬
coveries—Cleistogamous Flowers—The Part played
by Bees and Insects — The Banquet offered by the
Flower—The Parts of the Blossom—The Ovule and
Ovary—Embryo Sac — Formation of Embryonal
Vesicle — Pollen: its Structure — Pollen Tubes —•
Impregnation—Anatropous Ovule—Union of Nuclei
—Formation of Embryo and Seed—Monoecious and
Dioecious Plants—Aucuba japonica—Hazel Nut —■
Nectaries — Extra Floral Nectaries — Differences
between Nectar and Honey—Position of Nectary —
Nectar Cells: Microscopic Examination of; In
Pelargonium and Hyacinth—Aphide Honey.

The fact that bees gather both pollen and nectar
from blossoms has already been considered; but we
have yet to learn, why the wants of bees, in all their
genera and species, are supplied by the floral world.
The answer to this inquiry brings before us a new
meaning to the existence of all these insects, as
a part of that frame of nature in which nothing
is really isolated, although the bonds of mutual de¬
pendence may not always be apparent. Let us, first,

248

BEES AND BEE-KEEPING.

look at the matter in outline, filling in, hereafter,
such details as may seem necessary. Plants blossom
in order that seed may be produced and perfected,
and the race continued. But before seed, in the
true sense, can be produced at all, pollen, which is
borne by the anthers, and which we have all noticed,
as the abundant orange-coloured dust of the lily,
e.g., must be placed upon a certain special part of
the flower, called the stigma, a fact discovered by Sir
T. Millington two centuries ago. Should the pollen be
of a suitable kind, and the stigma in a receptive
condition, a delicate thread, called a pollen tube,
is thrown out, by the pollen granule, into the seed
vessel, by which the seed becomes fertilised, and,
when mature, capable of germination. The great
majority of flowers possess both anthers and stigmas.
They carry the two sexes within themselves ; and we
might suppose that, this being so, the form of the
flower would secure the transmission of its pollen
to its stigma, in order that the end of its being
might certainly be accomplished.

So thought the older botanists, and werejffn con¬
sequence, puzzled in explaining the reasons! for the
forms of the blossoms they examined. It wasMpointed
out, however, as long since as the close of the last
century, by a keen observer of Nature-—Sprengel—that
the structure of a large number of blossoms was such
as seemed designedly to render this simple arrange¬
ment impossible. His observations for many years
bore no fruit, and appeared to be overlooked; but,
during the last two decades, systematically-conducted
experiments and extended observations by many

BEES AND FLOWERS.

249

naturalists, especially Hildebrand, Hermann Muller,*
Delpino, and, above all, Charles Darwin,f have put
the whole question in a new light, and reduced
isolated facts to a law, which extends beyond the range
of botany. It is now shown that conspicuous flowers,
generally speaking, are especially modelled to prevent ,
or at least impede, fertilisation, by the pollen they
themselves produce ; while marvellous contrivances are
exhibited to secure pollen from some other plant or
flower of the same species ; for, amongst those that
have been studied in reference to this matter, there
exists but a very inconsiderable number of real or
apparent exceptions; whilst the latter, under renewed
examination, are not infrequently affording delight, as
they are found to possess some unsuspected adapta¬
tion to crew-fertilisation, which, in occasional instances,
especially amongst the orchids, | is so droll as to sound
rather like the outcome of a rampant fancy than a
narration of sober fact. I am not unmindful of the
existence of blossoms, denominated cleistogamous , pro¬
duced, under certain conditions, by some plants, and
which must, by their structure, be self-fertilised; these
we shall find, when they presently come before us, are
produced rather as supplementary or alternative than
exclusive organs of reproduction. The protest made
by Nature, for some profound, perhaps inscrutable
reason, against continuous in-breeding, applies, then, no
less to plants than to animals, to flowers than to bees.

* “ Die Befruchtung der Blumen durch Insecten,” 1873.

+ “The Effects of Cross and Self-Fertilisation,” and “The Different
Forms of Flowers on Plants of the Same Species,” 1877.

J “The Various Contrivances by which Orchids are Fertilised by
Insects.” C. Darwin. 1877.

250

BEES AND BEE-KEEPING.

But blossoms are fixed, if not even isolated. How
is the all-needful, fertilising dust to be carried from
one to the other ? For some, the work is done by
the wind, as when the blossoming corn is made to
gently rustle, or the lightly-suspended catkin of
the willow is vibrated in the upper boughs. Pollen,
in all such cases, having been formed, in countless mil¬
lions of granules, is, at its proper season, wafted by
every breath of air to the stigmas, made branched and
hairy to increase the chance of grasping it as it travels
past. But by far the greater number of flowering
plants confide to insects the duty of bringing about
those unions which, without them, would never be
effected. And, amongst insects, the whole family of
the Apidae are of the highest utility, followed by
butterflies and moths, while flies, and even humble
thrips, play their part; but it is the hive bee
especially that has been made the complement of
the blossom, the love messenger of the little beauties
of our woods and fields, supplying the eyes and
wings which have been denied to the flower itself.
As, then, the visits of insects are essential to the
existence of most plants, the flower secures these
by spreading a banquet, which it decorates with
its own beauty, and perfumes with its own sweet
breath. Pollen, it is true, is necessary for blossoms
themselves, but the amount produced is, without excep¬
tion, enormously greater than that required for mere
fertilisation; and the excess is the flesh-forming
food of the pollen-gatherer; while nectar—the basis
of honey, the heat and force-former, as grateful
to the insect palate as our own—is yielded, in the

FLOWER STRUCTURE.

251

great majority of instances, solely for her benefit.
Thus, then, insects perpetuate flowering plants, and
flowers continue the existence of insects, both being
but mutually sustaining parts of one great whole.

Let us now endeavour to follow the details by
which the general principles that have been sketched
are applied. If we take an ordinary flower—and, for
our present purpose, no better example can well be
suggested than the universal favourite, the common
geranium (pelargonium) of our gardens (Plate VIII.)—
and look at it from the outside, the first part brought
under our notice is a kind of cup—the calyx (c, A
and C)—here green, although in many flowers—the
fuchsia and larkspur, e.g .—it is coloured. Before the
blossom opens, when it is in the bud condition, this
cup incloses the internal parts, which are then in
the process of development, and protects them, in
their soft and tender condition, from external injury.
The calyx bursting as its contents develop, the

most conspicuous part of the flower, the corolla,
made up, in the pelargonium, of five, generally scarlet,
petals, begins to expand. The main function of the
corolla, in the greater number of blossoms, is to

attract insects, both by means of colour and scent.
Within the corolla we find the anthers (a), seven

in number, and differing altogether in shape and
appearance, both from the sepals—five of which

make up the calyx—and from the petals forming the
corolla. The calyx and corolla are of subordinate
importance, and may be regarded as protective and
decorative in character; but the anthers, which, to¬
gether with the stalks (the filaments) carrying them,

252

BEES AND BEE-KEEPING.

are called stamens, are absolutely necessary to secure
the reproduction of the plant. The anther is a double
bag, or, as in this instance, a pair of such, containing
a quantity of tiny granules (pollen), which, in reality,
are individually highly organised parts, capable, as
already hinted, of bringing about the wondrous pro¬
cess of fertilisation. Lastly, within the space lying
between the encircling stamens, and occupying always
the centre of the flower, lie the female reproductive
organs (s, C), collectively denominated the pistil.
This assumes very diverse characters in different
blossoms; but here, as in all the more perfect forms,
it consists of three parts, the bottom one being a
pouched cavity ( o , B), or hollow receptacle, called
the ovary, because it contains one or many minute
egg-like bodies (the ovules), each inclosing a germ
cell awaiting impregnation. Rising from the apex
of the ovary is a stalk-like part (the style), seen
beneath s, C, and surmounted at its summit by a body
of peculiar structure, called the stigma, which, at a
certain point in its development, becomes capable
of receiving the pollen granules—really the homo-
logues of the sperm 'cells of the drone (page
216). The flower, as such, in all cases is a means,
not an end, and, as the latter is accomplished, it
disappears. The corolla, having caught the insect’s
eye, dries; the stamens, having yielded their sperm,
wither. The stigma and the style, having performed
their office, dry and shrivel, and nothing is left
except the ovary, which is, in some cases, surrounded
by a persistent calyx. The ovary now grows and
develops into what is called the seed vessel, but

FERTILISATION.

253

botanically, the fruit ; here, at length, we have the
seeds, which are the ripened ovules, while the seed
vessel is the ripened ovary. But the change from
ovule into seed is not merely one of growth; it
depends upon the formation within the ovule of the

Fig. 49.—Ovaries, Ovules, Pollen Grains, and Tubes.

,, Section through Ovary of Buckwheat (Polygonum fagopyrum)—s, Stigmatic
Surface; pa. Pollen Grains; st, Style; pt, Pollen Tube; ov, Ovule ; s, Secun-
dine; p, Primine; n, Nucellus; es, Embryo Sac ; emv, Embryonal Vesicle.

B, Section of Pistil of Pansy (Viola tricolor)—s, Stigma; l. Lip; pt, Pollen
Tube; ov. Ovules. C, Pollen Grains Various, Emitting Tubes— s, Portion of
Papillose Stigma. D, Ovule at a very Early Stage— n, Nucellus; es. Embryo
Sac fonr 5 "" • *» Jm *- n 'PrlminA • s liittn Rficnnduift. TS. Tiftvelfined 1 ■

Ovule oi

embryo, which is itself the outcome of the definite
process of fertilisation, now to be examined in detail.
Taking a blossom of buckwheat, well-known as an

254

BEF.S AND BEE-KEEPING.

abundant honey-plant, removing the corolla, and making
a section through one of the ovaries, we find, within a
cavity, the ovule ( ov , A, Fig. 49), which, on account
of its being straight instead of curved, and solitary
instead of one of a number, is an excellent subject
for study, and, for the present, may serve as a
type of those formed by flowering plants in general.
When examined minutely, it is observed to consist
almost entirely of cellular tissue— i.e., of a number
of minute sacs, similar to those at G (Plate VIII.),
placed side by side, in close juxtaposition, forming
( n , A, Fig. 49) the nucellus of the ovule, which
is enveloped in two coatings of firmer texture, called
primine ( p) and secundine (j), which grow up over
its surface in its early days, as shown at D. These
surrounding layers, however, are. never continuous
over the apex of the ovule, where they leave an
open channel, called the micropyle, quite similar in
function to the micropyle of the egg (page 231),
and here permitting communication with the nucellus,
and a large cavity within it, called the embryo sac
(es). In the vast majority of plants, the two sides
of the ovule are unequally developed, so that, during
its growth, it is made to turn partly or completely
over, as in the anatropous ovule (E), where the
arrow indicates the micropylar aperture, or in the
ovules of Viola tricolor ( ov , B), one of which is
shown more enlarged at F.

The extremely interesting and instructive history
of the formation and development of the embryo
sac, with the mystic and involved movements which
prepare for and accompany fertilisation, can only

FERTILISATION.

255

here be touched upon. A cell near the apex of
the nucellus undergoes division, which is repeated
until a line of cells, with thick Avails, has been
formed. The lowest cell of the number now
enlarges greatly, at the expense of the others,
which are absorbed, and an enormous cell (the em¬
bryo sac, es, A and F) results. During the growth of
the latter, its nucleus divides, and the two new
nuclei travel to its opposite ends, a large central

Fig. 50.—Embryo Sac, before Fertilisation, in Three Stages op
Development.

A, B, and C— h, Helper Cells; a, Antipodal Cells, derived from Division of
Original Cell Nucleus ; pn, Polar Nuclei ; dn, Definitive Nucleus ; em.v,
Embryonal Vesicle.

sap cavity, called a vacuole,* being formed. These
nuclei, now stationed at the upper and lower ex¬
tremities of the embryo sac, twice divide into two,

* The life of the cell inheres in its protoplasm. When this separates
and gives place to cell sap, the spot occupied by the cell sap is
called a vacuole.

256

BEES AND BEE-KEEPING.

four nuclei resulting in each case. Mysteriously, one
nucleus starts from each end (/>«, pn, A, Fig. 50),
approaches ( pn , B), and at last meets, its companion
in the centre, and coalesces with it to form the
definitive nucleus ( dn , C). These two nuclei are
called the polar nuclei. Round the two sets of
the three remaining nuclei a process of free cell
formation begins, resulting in three cells at each
end of the sac instead of three nuclei. Those at the
lower end (a, B and C) soon become surrounded
with cell walls, while those at the micropylar (upper)
extremity remain naked, and constitute the egg
apparatus. Two of these lie above the third, the
latter constituting the oosphere, or embryonal vesicle
( em.v), which has its nucleus lying at its lower
end. Generally, all three cells of the egg appa¬
ratus position themselves in contact with the wall
of the embryo sac, which is, at this time, awaiting
fertilisation from the pollen previously placed on the
stigma, it may be by wind action, by the gardener,
or by the little professional pollen-carrier, the nature of
whose burden we must now endeavour to understand.

Pollen granules vary greatly in form, colour, and
size. They are frequently approximately spherical,
sometimes oval, triangular in the fuchsia or evening
primrose, hexagonal in the chicory, covered with
minute spines in the hollyhock and aster, curiously
banded in the Passion flower, spirally grooved in
the musk ; while they are, in these and other similar
plants, delicately coated with an oily body, giving them
adhesiveness, and aiding the bee in packing them upon
the legs. When cut into sections, they reveal a complex

A, Pelargonium Blossom— s, Stigma ; c, Calyx ; n, Nectary. B, Calyx, with Ovary in
Cross Section— n. Nectary ; o, Ovary. C, Blossom, side view (Corolla removed)—
•s, Stigma; a, Anther; c. Calyx. D and E, Cross Sections of Ovary through
lines a and b of A. F, Stamen—a, Anther. G, part of E (Magnified 180
times)— cm, Cuticle; gh, Glandular Hair; h, Hair; nc, Nectar Cells. H,
Longitudinal Section through Upper Part of Nectaiy— gh, Glandular Hairs.
I, Longitudinal Section through Lower Part of Nectary—nc, Nectar Cells. If
and L, Sections of Nectar Cells (Magnified 500 times)— n, Nectariferous Nucleus.

FERTILISATION.

257

structure: hollow within, they are filled with a very
granulous protoplasm, with many oil globules ; their
solid coat is formed in two layers, an inner and an
outer, called intine and extine. The pollen grains
are developed within the anther, by constant seg¬
mentation of the contents of the latter. It appears
that, when the grains become isolated from each
other, the nucleus of each one divides into two un¬
equal parts, the smaller of which attaches itself to the
wall of the granule. When the pollen grain is placed
upon the moist stigma, so that nourishment is given
to it, the interior parts grow, and burst the exterior
enveloping coat (the extine), at points where it is
curiously thinned down, while the intine is corre¬
spondingly thickened—the blunt angles of the pollen
of epilobium, e.g. (D, Fig. 57), have a delicate pellicle
of extine only, but here the intine ( ti) is extremely
strong and dense ; the latter, therefore, remains un¬
ruptured, and holds the protruding interior, elastically
extending with it (E), so that, under favourable con¬
ditions, a tube of extraordinary length is developed,
through which the larger nucleus at last passes.
Pollen grains, placed in soda-water sweetened with
a little sugar or honey, will, if kept in a genial
temperature, grow under the microscope, giving some
such forms as shown at C, Fig. 49. They may often
be found in the stomach of the bee with a tube
partly developed, while in the later food of the
larvae this phenomenon is quite common.

During the curious sequences represented by
A, B, C, Fig. 50, the stigmas, usually covered by
little papillose bodies, coat their surfaces by secretion

258

BEES AND BEE-KEEPING.

of a sugary, glutinous fluid, which causes the pollen
grains to adhere ; the pollen tubes, which seem to re¬
ceive nutrition as they push forward, now penetrate,
with astonishing rapidity, and to surprising distances,*
either passing through the channel of the style (pt, B,
Fig- 49 )> or its loose conducting tissue (st, A), into
the cavity of the ovary, where, in the darkness, they
travel on, as though endued with intelligence, un¬
erringly finding the apertures in the primine and
secundine (the micropyle), by which one enters and
applies its now swelling end to the extremity of the
embryo sac. The two upper cells of the egg appa¬
ratus ( h, C, Fig. 50) in some cases absorb the end
of the wall of the embryo sac ; but always—bv
methods, subject to variations in different orders—the
pollen tube transfers its protoplasm and nucleus,
by the agency of these helper cells, to the embryonal
vesicle. Since every ovule needs a pollen tube to
fertilise it, the number of tubes requisite will depend
upon the number of ovules, but usually many more
are produced than can be utilised. In the buckwheat,
e -S- ( A , Fig- 49 ), we find but one ovule in each
ovary, while in many plants, especially orchids, they
are multitudinous. At B (the ovary of the pansy), six
are represented, but many more actually exist; and
here we notice how beautifully suited to the exi¬
gencies of fertilisation is the turning of the ovule
by unequal lateral development. Had these six
ovules stood straightly up, like that of the buck¬
wheat, their micropylar apertures would have been
placed in an exceedingly unfavourable position for

* In the common crocus, the style is frequently several inches in length.

FERTILISATION.

259

meeting the entering tubes; but the tiny cavity is
turned round from the centre, to face the wall of
the ovary (F), which is slightly hairy within.
Clinging to this hairy surface, the tubes feel their
way along, to find, and at once enter, the point they
seek. The helper cells z, Fig. 50) now disappear,
while the embryonal vesicle becomes granular, and
two nuclei can be detected in it. One of these is
the nucleus of the oosphere, or embryonal vesicle;
the substance of the other has, doubtless (according
to Sachs), been derived, through the helper cells,
from the pollen tube. These two nuclei, male and
female in their origin, meet and coalesce, constituting
the nucleus of the fertilised embryo, or new indi¬
vidual, which now surrounds itself with a cellulose wall,
and so starts an existence, which yet depends upon
nurture derived from the female parts of the parent
flower. When it has acquired some development,
and a supply of food sufficient to enable it to initiate
a separate existence, it will be cast off as a
mature seed.

We have, up to this point, spoken of flowers as
though they invariably carried both stamen and pistil,
and usually this is the case; but exceptions are not
infrequent. Every one knows that, in the melon,
vegetable marrow, cucumber, and other plants belong¬
ing to the order Cucurbitaceae, some of the flowers
are male, while others are female, the latter bearing
the fruits. In these cases, it is obvious that the pollen
necessary for the fertilisation of the ovule must be
carried, by some means, from one form of flower to
the other; and when, by the method of culture, insects

v 2

26 o

BEES AND BEE-KEEPING.

are excluded, the operation denominated “fertilising,”
or “ setting,” is undertaken by the gardener. The two
genders of unisexual flowers, sometimes placed, as in
the vegetable marrow, on different parts of the same
plant, hence called monoecious , are frequently produced
on distinct plants—then called dioecious , meaning two
houses which are, necessarily, the complements to
one another. This fact was known to Herodotus, in
the fifth century before Christianity, who describes
the process of “ caprification ”—the transference of
pollen from the male blossoms of one tree to the
female blossoms of another—by -which a crop of dates
was insured on the Egyptian palms.

In our own day, a curious instance has occurred,
in the case of the Aucuba japonica (the common
blotched laurel), a single plant of which was long
since introduced into this country by the Dutch.
This solitary specimen, from which, up to a few
years since, all the countless plants decorating our
gardens and shrubberies had been derived, by cuttings,
happened to be a female. The myriads of ovules
formed in all the inconspicuous, chocolate-coloured
flowers of the descendants of this parent, of course
invariably withered for want of fertilisation; but,
a few years since, the male Aucuba reached us,
and beautiful scarlet berries began to be formed,
and our ancient friend made additionally attractive
as a decorative plant. These berries gave us new
individuals, exhibiting variations from the parents,
and yielding some male, some female, flowers, so
that the berrying of the laurel is already general.

The common hazel bears unisexual flowers, which are

DICECIOUS PLANTS.

261

utterly dissimilar ; the males [a, Fig. 51) are grouped in
drooping lines, called catkins, each containing some¬
thing more than a hundred flowers, which come out
soon after Christmas, remain on the tree for a few
weeks, and then drop. But they have accomplished
their work, for the ten or twelve anthers each blossom
carries furnish abundant pollen, which, shaken by every
breeze, and being non-adhesive, gently falls through
the spreading branches below, where we may find

Fig. 51.—Inflorescence of Nut.

a, Staminate (Male) Catkin of Nut; b, Pistillate (Female) Blossom of ditto
c, Pistillate Blossom, Unopened.

small, hardly observable female flowers (d), consisting
of hairy, branched stigmas, crimson in colour, and
rising from amidst a few small scales, which con¬
ceal the ovary. The stigmas catch the dropping
granules, the pollen tube is thrown out, and fertili¬
sation follows, preceding, in order of time, the
expanding of the leaves, which would, if opened,
seriously impede the operation. The necessary
abundance of pollen, since so much is inevitably

262

BEES AND BEE-KEEPING.

wasted, gives an excess, of which the bees take
advantage ; and often, in the early spring, the stocks
are greatly helped by the catkins, not only of the
hazel and other nuts, but also of the beech, the
poplar, and the willow (of which the two catkins
are represented in Fig. 52). It is curious that, in
the case of the weeping willow, notwithstanding its
wide distribution, only pistillate (female) trees are in

Fig. 52.—Catkins of White Willow (Salix alba).

A, Staminate (Male) Catkin; B, Pistillate (Female) Catkin.

cultivation, which must have all originated from a
single parent. The blossoms just mentioned are wind-
fertilised (except the willow, whose position is inter¬
mediate), and form a few examples of those called
anemophilous ; while those depending on insects are
denominated entomophilous.

Before proceeding to examine the various remark¬
able modifications made in flowers, in order that the
pollen produced by their anthers, in the closest proxi-

NECTARIES.

263

mity to the stigma, should yet not fertilise the latter,
we must discuss the manner in which the nectar is
produced, and placed so as not only to attract the
insect, but also force it, while taking its repast, to
deposit pollen, brought upon its body, on to the
stigma.

It is more convenient than accurate to speak of
“honey-yielding plants,” and of bees gathering honey;
for the fluid secreted by the flower is unlike honey
in more particulars than one, and is denominated
nectar, while the part by which it is yielded is called
a nectary. Although it is certain that the character
of the secretion varies considerably in different plants,
analysis has shown that, in a large proportion of
instances, the sugar it contains is identical with that
derived from the cane or beet-root, while the sugar
of honey is similar to that of the grape. From what
has already been said of the glandular and tongue
structures of bees (pages 81 and 101), it is clear
that a salivary secretion is added to the gathered
nectar, and that this, like the saliva in our own case,
converts the cane into grape sugar; and probably
also, as with ourselves, this is an initial step in
assimilation, since cane sugar is actually poisonous
to the blood, while grape sugar acts within it as a
normal producer of heat and force. Many flowers
are especially contrived for fertilisation by moths
and butterflies, and there is strong reason for sup¬
posing that these latter insects produce exactly the
same alteration—technically, “ inversion ” of the sugar
of nectar—as our bees.

From what we know of the chemical changes occur-

264

BEES AND BEE-KEEPING.

ring during the germination of seeds, and in leaves
when stimulated by light, we should expect sugar to
be present in considerable quantities in flowers, where
growth is so rapid, and cell energy so apparent.
Many careful observations, made of late years, by
botanists, in various countries, have shown, amongst
other interesting facts connected with the existence
of nectar in plants, that flowers contain it in quantity,
in their tissues, even when no nectary is present to
secrete it; and also that, in vegetative organs, quite
apart from the inflorescence, nectaries are occasionally
present— eg., in the bracken fern (Pteris aquilina ),
nectar flows from small, pale swellings at the bases
of the secondary petioles; and the stipules (or leaflets
on the leaf-stalk) of beans are nectariferous, as are
also small glandular prominences on the leaf-stalk of
a species of Prunus , and little, brownish pittings in
the leaf-blade of some laurels. From the latter I have
sometimes seen hive bees gathering industriously,
while their visits to bean stipules are quite en regie.
It is here very interesting, while practically important,
to note, that experiment has shown that emission of
water vapour into the atmosphere, and emission of
nectar on the surface of the nectary, are so related, that
what favours the one retards the other, the damaging
effect of a prevailing east wind being thus perfectly
explained. In the flowers, nectar is usually furnished
most abundantly in the early morning, diminished
till afternoon, and again increased towards evening.
Although high temperature favours secretion, flowers
of the same kind yield larger amounts in colder than
in warmer climates.

NECTARIES.

265

To trace out the probable development of the
nectary is beyond our limits; but, in a word, if it
be granted—and experiment is conclusive on the
point—that intercrossing does lead to greater vigour
in the resulting seed, then any variation making
intercrossing more certain will lead to a selection
favourable to the individual presenting that variation ;
so that transudation of sugary matters forming a
rudimentary nectary, and so attracting insects, will
tend to establish and extend the variation, by which
the flowers will become permanently nectariferous.

The position of the nectaries in flowers, and the
organs of which they are modifications, differ with
the kinds of insects for which they are suited; some
lie almost on the surface of the flower— e.g ., in the
carrot, elder, ivy, &c.—but most are situated in its
deeper recesses, not only because this position draws
the visiting insect well into contact with the male
and female parts, but also because exposure to water,
in the form of rain or dew, injures the nectar, and
decreases its attractiveness. This fact is the counter¬
part of the enormous length of proboscis possessed
by moths, butterflies, and bees. In many flowers,
strange devices save the sugary fluid, even in the
most persistent downpour— e.g., in the upstanding
white dead nettle, the upper lip is formed into an
umbrella (see Fig. 66) ; in the Tropaeolum majus
(garden nasturtium), upright water-resisting hairs (see
Fig. 55) prevent rain travelling towards the spur ; and
in the useful Borago officinalis (borage), the drooping
habit of the flower, and the tube-like cavity formed
between the stamens, give perfect protection.

266

BEES AND BEE-KEEPING.

The fact that sugar is present in flowers because
of their rapid development, would lead us to expect
it in greatest excess where the energy of life is
most intense—and this is in the ovary ; and, sugges¬
tively, it is in this neighbourhood that the nectary is
far most commonly found, numerous instances presently
coming before us. Curious variations, however, occur.
Poplars, which are anemophilous and dioecious, yield
so much sugary secretion on the stigma, whose office
it is to glue down the pollen granules floating by,
that the stigma really becomes a nectary; and these
trees, although altogether independent of insect action,

Fig. 53.—A, Appendage of Anther, forming Nectary of Viola tricolor
(Pansy), Order Violacece (Magnified Twenty-four times)—me, Nectar Cells,
exaggerated. B, Anther as Removed from Flower— a, Anther Cell;
nc, Nectar Cells.

yet yield a restricted quantity of honey : in some, the
base of the style, in others, aborted stamens, become
nectar-yielding ; the transuding syrupy surfaces often
appear on the petals, as in some species of butter¬
cups, where they are covered with a small, flat scale,
behind which the nectar is formed; or on the sepals,
as in the lime, so well loved by bees; in many,
the petals are rolled into a tube, as in the colum¬
bine, hellebore, aconite, &c., and the inner end of
the organ is the nectary; but in some— e.g., the
violet—the spur merely serves to receive the nectar.
In Viola tricolor (the pansy), the larger and lower

NECTARIES.

267

petal is thus extended backwards, and curious ap¬
pendages ( nc, B, Fig. 53) on two anthers pass into
the cavity provided, and there secrete a sweetish fluid.
Perhaps no flower presents equal advantages with
this to the microscopic tyro who would study the
structure of the nectary, and the cells (nectar cells)
which yield the secretion ; for not only are these large
( nc , A), characteristically sugar-loaf-shaped, and promi¬
nent, but they lie on the outside of the process (their
protection being derived from the covering afforded
by the spur-like petal previously mentioned), and,
consequently, the difficulties of section cutting are,
in this case, altogether avoided. The nectar-producing
tissue is usually made up of small, thin-walled cells,
containing abundant protoplasm, a nucleus, and cell
sap, rich in sugar. Often the nectary shows a number
of pores, or stomata, on the surface-layer of the cells
which line it, and through these the nectar is poured
on to the face of the organ, whence it may be
sucked up by the visitors to the flowers. Where
pores are absent, the covering membrane is extremely
delicate, permitting either free transudation, or yield¬
ing at once, as in some orchids, to the abrading
action of the insect tongue.

Returning to our pelargonium (Plate VIII.), selected
because it is at command in most places, and at
every season of the year, we find, running down the
flower stalk, and immediately under the uppermost
and broadest sepal, an enlargement of the stalk itself,
marked off by inconspicuous grooves, and terminating
in a small bulbous expansion a little below the line
b (A), and which is often purplish in colour. This is

268

BEES AND BEE-KEEPING.

the nectary, and is really formed by carrying the
upper part of the calyx down the stalk. If we com¬
pare its position and relations to the rest of the
flower with those of the Tropaeolum majus (Fig. 66),
which is of the same order ( Geraniaceae ), we shall
find that the difference lies in the latter nectary being
free, while the former is attached (adnate) to the
pedicel, or stalk, of the flower. If we now remove
the petals, and look at the calyx from the front, we
see into its opening [n, B). Making cross sections
through the lines a and b, we find the nectary wider
above, as at D, and narrow below, as at E. A keen
razor, dipped into methylated spirit, will take off slices
sufficiently thin for microscopic examination, under
a cover glass, in water. Cutting D longitudinally,
so that the nectary is divided, and then removing a
thin slice from that which forms the upper part of
the Figure, and magnifying about 200 diameters, we
find the outside to consist of cuticular cells, carrying
glandular hairs (gh, H), which secrete a resinous
body, of strong odour. The cells on the opposite
side of the section are not unlike those of the ex¬
ternal cuticle, because they have here no secretory
function, although they constitute the lining of the
upper part of the nectary. Taking a section (G)
from the face of E, which lies in the line b (A),
we discover the hairs and cuticle to be of precisely
the same character as those previously noticed; but
the lining cells ( nc ) of this part of the nectary are
totally different, extending inwards by almost pointed
prominences. Now cutting E longitudinally, and
taking from it a thin section, we find the lining cells

NECTARIES.

269

all pointed, as at I, where they face those which lie
near the mouth of the nectary, as a part of H. The
structure of the pointed cells is quite special, their
contents, as seen under high magnifying powers,
being distinctly granular, peculiarly so near the cell-
wall, which, at the prominence, is excessively thin,
and has, lying immediately within it, a globular mass
of highly refractive protoplasm (n, K and L), con¬
taining a distinct nucleus. This is the active agent
in accomplishing the secretive act, and the surface of
the cells here, in healthy plants, and in proper con¬
ditions of the atmosphere, will always be found to
be coated with a layer of nectar.

We may study, similarly, the three nectaries of the
common hyacinth If the corolla be removed, we
find the flask-shaped ovary giving indications of being
formed by the fusion of three parts. The furrows
running between these carry, near their upper ends,
tiny beads of nectar, secreted from a tube-like cavity,
running down between the cells of the ovary; and, by
making cross sections, we get an opportunity of
examining the nectar cells. But, in some cases, we
find no superficial layer possessing the secretory
function, but an alteration in the underlying cellular
tissue, which carries its nectar onwards to a pore
passing through the ordinary epidermal cells. This
structure obtains in the raspberry (Fig. 70), where a
continuous line of pores (no), commonly covered by
beadlets of nectar, easily seen by a lens, surrounds the
drupels; the nectar being, of course, secreted by
nectar cells ( nc ), which are not superficial, but form a
part of the receptacle of the blossom.

270

BEES AND BEE-KEEPING.

It would be well for apiculture if plants yielded to
bees no other nectar than that flowing from their
blossoms ; but, unhappily, a sort of second-hand honey,
primarily derived from plant juices, and of very
objectionable quality, is frequently gathered from
the insect pest, the Aphis, or plant louse, in such
quantity as to utterly ruin the legitimate harvest.
The whole question of this pseudo-honey has very
great interest, and demands the careful attention both
of the gardener and bee-keeper.

None can have failed to have noticed the shining
and gummed appearance frequently presented by the
leaves of the lime, the sycamore, the oak, the maple,
and the elder, particularly in hot weather; while the
plum, the apple, rose, and currant—amongst many
other plants and trees — are often brought into an
almost disgusting condition from the glutinous liquid
which covers them, and which, because anciently sup¬
posed to be a deposition from the atmosphere, re¬
ceived and retains the name of “ honey dew.” Kirby
and Spence, in their “ Introduction to Entomology,”
say: “You have, doubtless, observed what is called
the honey dew, upon the maple and other trees, con¬
cerning which the learned Roman naturalist, Pliny,
gravely hesitates whether he shall call it, the sweat
of the heavens, the saliva of the stars, or a liquid pro¬
duced by the purgation of the air. Perhaps you may
be aware that it is a secretion of Aphides, whose
excrement has the privilege of emulating sugar and
honey in sweetness and purity.” De gustibus dis-
putandum est, and certainly here but few would en¬
dorse the closing words of these authors. Plants

APHIDE HONEY.

2 7 I

rarely, and probably only in diseased conditions, se¬
crete excessive quantities of sweet liquid, which, oozing
from various parts of their surfaces, gives the eager
gatherer material that is above suspicion; but ordi¬
nary honey dew is now universally conceded to be

Fig. 54.—Aphides as Nectar-producers.

A, Rose Aphis ( Siphonophora Rosce), Winged Viviparous Female (Magnified Ten
times)— an, Antenna; ; n, Nectaries ; r, Rostrum, or Proboscis. B, Wingless
Oviparous Female of same— an, Antennae; n, Nectaries ; ng, Nectar Globule;
e, Egg just laid. C, Aphis Scabiosce (Aphis of Scabiosa arvemis). Wingless
Viviparous Female (Magnified Sixteen times) — an, Antennae; n, Nectaries;
dy, Developing Young ; y, Young Aphis, just born. D, Aphis Sambuci (Aphis
of Elder Tree), Wingless Viviparous Female, with Ant Feeding on (gig)
Nectar Globule— an. Antenna; ; n, Nectaries; an', Antennae of Ant.

the product of the Aphis. When the gummed leaves
are lifted, they will be found to be infested beneath
by colonies of these creatures, some winged and some
wingless; and a careful examination will generally

272

BEES AND BEE-KEEPING.

show that they are provided with two short tubes,
called the nectaries ( n , A, Fig. 54), by which they
are enabled to eject a sweet fluid.

Leaves are constantly forming starch, which is at
once converted into the soluble form, sugar; so that
the Aphis is, perhaps, provided with saccharine sub¬
stances in such quantity that the excess must be
drained off. Standing, two or three years since, in the
shadow of a lime tree, I saw falling, in the sunlight,
a thick, constant shower of minute drops, which were
being expelled from the anal apertures and nectaries
of the Aphides infesting the leaves. The necessity
for this vigorous ejection is apparent; without it, the
closely-packed colonies would soon be hopelessly
fixed to the leaf, and to one another. The grass be¬
neath the tree was thickly gummed, while the upper
surface of every leaf was closely covered, and not
a few bore incipient drops at their points. Other
instances quite as remarkable have attracted my
attention ; in one case, that of a sycamore, overhanging
some paving-stones, the latter were rendered actually
dangerous to the pedestrian ; and one of my apple
trees (“Sturmer Pippin”), this summer, became, in very
few days, so covered, that every leaf carried hundreds
of the Aphis Mali , and every fruit was running with
what might have passed as a compound of treacle and
soot. Few botanical families appear to be altogether
proof against the attack of these pests, which are all
but universally distributed, and of which Beckton,* in
his magnificent work, describes about 300 distinct
species, the oak suffering from about six, the birch,
* “ Monograph of the British Aphides,” Ray Society, 1883.

APHIDE HONEY.

273

willow, and fir, from eight each, the elm from four,
and the currant bush from three.

Their rapid multiplication and very injurious effect
cease to be a wonder when we learn something of
their habits and capabilities. The male only appears
amongst them at intervals, which may be distant, and
he has only been well made out by dissection in
about twelve species. An impregnated egg having
been deposited, very many generations of individuals,
formed by a process of interior budding, and born
alive, will succeed one another, before the cycle is
completed, and the fully-sexed female and male give
again origin to the impregnated egg. Usually, some
days after the appearance of the male, the oviparous
female (B, Fig. 54) begins to deposit her eggs,
which, in most species, are relatively enormous, each
one ( e ) equalling in length half the body of the
mother. When deposited, they are coated over with
a glairy fluid, attaching them to twigs or stipules ;
they are then pale, but soon become brown or black,
and are capable of bearing the most intense cold of
winter. Nor are the Aphides themselves much less
hardy. Beckton states that he witnessed the hatching
of a young Aphis from the eggs of Siphonophora Rosse
(the Rose Aphis), on March 12, 1873, when the ther¬
mometer stood at 25 0 , and most species can endure
lower temperatures without visible injury. The insect,
after leaving the egg, very rapidly grows, and quickly
attains its full size, exhibiting in its pseudovaries (false
ovaries) developing larvae, which soon begin to make
their escape at the rate of many daily. Indeed, the
body cavity of the viviparous Aphis, during the summer

274

BEES AND BEE-KEEPING.

time, is almost exclusively occupied by the embryos
(dy, C, Fig. 54) and the digestive apparatus. If an
adult female be removed from the under side of the
leaf of a rose bush, and the abdomen snipped, as many
as thirty immature Aphides may often be seen to escape,
by applying a little pressure, under the microscope.
The embryos are in all stages of development, those
lying nearest the body aperture being the largest, and
showing the eyes, antennae, and limbs fully formed.
The description previously given of the ovaries of the
queen bee (page 213) will aid in understanding the
ovarian chamber in this smaller insect. They are
gathered into tubes, which are, again, formed into two
bundles, disposed laterally, each communicating with
its own oviduct. The upper extremities of the ovarian
tubes are very attenuated, and lead into a chamber
where the germinal matter is elaborated. Here
nucleated cells are visible, which, according to Brandt,*
have an amoeboid, movable nucleus, and correspond
to the ordinary germinal vesicles, with their usual spot ;
the ovum becomes the larva, which is extruded fully
formed, as we have already seen. The progeny, even
at the time they quit the parent, show the traces of
another generation within themselves ; thus, a single
insect, hatched from one of the shining black ova,
may, during her lifetime, be the mother of many
billions of young. Reaumur calculated that one
Aphis may give origin to the enormous number of
5,904,900,000 individuals during the month or six
weeks of her existence. Happily, Aphides have many
insect enemies, which, as our friends and helpers, must

* “ Ueber das Ei und seine Bildungsstatte,’’ 1878.

APHIDE HONEY.

275

receive attention before we close, while rough weather
plays havoc with them, rainstorms sweeping them
away by myriads ; but, upon the supposition that no
casualties occur, Reaumur’s figures are far too low,
and Tougard and Morren show that a quintillion are
within the efforts of a single mother ; and Professor
Huxley* gives the amusing calculation that, assuming
an Aphis to weigh jo^-g-gr., and a man 2851b.— i.e.,
2,000,000 grains—then, the tenth brood of one parent,
without adding the products of all the generations
which precede the tenth, would contain more ponder¬
able substance than 500,000,000 of such men— i.e .,
more than the whole population of China.

Entomologists formerly thought that the production
of the male which consorted with the female was
brought about in anticipation of the close of the
season, so that the wintering egg might be produced ;
but this does not appear to be accurate, and recent
observations show that viviparous reproduction may be
continued during several seasons, but that, at length,
recourse to a new infusion of vitality by ordinary
sexual means becomes necessary. A scarcity of food
tends to the formation of winged females (A), capable
of repairing to new pastures. These are invariably
viviparous, while the wingless females may be either
oviparous (B) or viviparous (C), from which it is
noticeable that the larva is expelled tail foremost.

Ants are particularly fond of the sweetish exudation
of the nectary, and they frequent the haunts of the
Aphis, beating on the sides of the insects with their
antennae, when the liquid is at once driven out, as
* Huxley “ On Organic Reproduction of Aphis.”

Z 2

276

BEES AND BEE-KEEPING.

seen at ng, B and D, the ant in the latter figure greedily
swallowing what the Aphis offers. Linnaeus, in con¬
sequence, called the latter insect the cow of the ant
(“ Aphis formicarum vacca ”), and Darwin and Sir John
Lubbock, amongst others, have shown that the ants
almost literally milk the Aphides, which seem to
attempt to retain the secretion until the ants are
ready to receive it, of which they give indication as
just now noted. The demand, unfortunately, is in no
way equal to the supply, and so this aphide honey is
thrown out, to fall on to the upper surface of the leaves,
where it is gathered by bees, especially after rain, which
renders it sufficiently liquid to permit of its ready
removal; but its taste is mawkish, its odour not pleasant,
and its colour often as dark as treacle, and of a dirty
hue. That gathered from the sycamore and oak is ex¬
tremely black, and ought not, in my opinion, to be
regarded as fit for human consumption.

All are interested in reducing the numbers of this
obnoxious insect, whose fecundity is so prodigious ;
but, after all the schemes that have been propounded
for its destruction, it seems evident that we must look
to Nature’s own checks, aided by any encouragement
or protection that we may be able to give to Aphis-
devouring creatures. Foremost amongst these come
the numerous species of Coccinella (Ladybird), the
food of which consists almost exclusively of Aphides.
Their marvellous voracity is shown equally in their
larval and winged condition. In the former stage,
the colour is slaty grey or brown, while the body is
covered with tufted tubercles, and provided with
mandibles, efficient both for holding and sucking out

APHIDE HONEY. 277

the juices of the prey, which is seized by the back,
and the liquid contents quietly sucked out, the whole
process requiring about a minute. Ladybirds, if dis¬
covered clustered in crannies, in winter—and they
sometimes collect many thousands together—should on
no account be destroyed.

Some of the most familiar and the most beautifully
coloured flies of our summer belong to the family
Syrphidas , which, from the peculiar character of their
hovering, darting flight, have been popularly called
“ Hoverers,” and the larvae of several species of these
devour Aphides in immense numbers. These creatures
are legless, blind, and leech-like in form, and move
slowly, by means of hooklets, with which the posterior
rings of their bodies are furnished. The maggot,
after each advance, makes a lashing motion with its
head, in search of food, and, when an Aphis is struck,
it is taken off its legs, and hoisted into the air, where
its juices are extracted, and the skin rejected. The
eggs of these flies may often be found deposited in
the midst of Aphides, multiplying to provide food for
the larva at the time of its hatching.

The beautiful Lacewing flies, whose green bodies,
delicate wings, and golden or red-tinsel eyes, are so
universally admired, are, in the larval state, great
enemies to Aphides. When fully fed, the larvae attach
themselves to leaves or stems, and change into short,
oval pupae, hanging head dowmwards. Amongst the
smaller Hymenoptera are found very many most use¬
ful destroyers of Aphides— e.g., the Cynipidx, although
usually regarded as injurious to many plants, as
gall makers are, in some of their species, serviceable,

278

BEES AND BEE-KEEPING.

depositing eggs in the Aphis of the rose, the parsnip,
the willow, the plum, the peach, and many others.
We may also reckon as allies many tiny Ichneumon
flies, which persistently attack Aphides not much
smaller than themselves. The female deposits from
one to five eggs within the body of the Aphis. The
resulting grubs live on the food assimilated by the
host, whose vital organs, with much consideration,
they do not invade until the last moment. His ex¬
ternal skin they leave, as this forms a case, within
which they pass through their pupal change. Aphi-
dius Rosas, Beckton tells us, he watched “ for some
time, as two individuals seated themselves upon the
backs of Aphides, seeming to enjoy the contortions
made to throw them off. After about five minutes,
the ovipositor was inserted by a sort of thrust, when
the flies made away in pursuit of other game.” As
the grub within develops, the Aphis visibly undergoes
an abnormal modification, the skin at last hardening
into a globular, horny box, within which the perfect
parasitic Ichneumon is formed, and at last escapes
by carving a circular hole. These cases, pierced and
empty, may constantly be seen under rose leaves.
Curiously enough, the above-mentioned changes are
often interrupted by a second parasitic attack, another
egg or e SS s being deposited within the first parasitic
larva, so that it, in turn, succumbs, and another form
of life presents itself. How endless the interchanges
of life and death ; respecting which the philosopher is
not much in advance of the humorist, who says—

“ Larger fleas have lesser fleas upon their backs to bite ’em,

And these, again, have smaller fleas, and so ad infinitum

CHAPTER XVI.

BEES AS FERTILISERS, FLORISTS, AND FRUIT-
PRODUCERS.

Method of Study — Tropaeolum majus — Anthers:
Movements of — Colour Streaks on Petals : their
Uses — Delphinium — Scrophularia nodosa — Water
Ejected by Bees—Epilobium angustifolium—Kalmia
latifolia -—Poisonous Honey—The Retreat of the
Ten Thousand — Heather—Erica tetralix—Calluna
—Erica Unedo — Berberis — Dimorphism — Primula
veris—Linum grandiflorum — Trimorphism—Lythrum
sal ica ria—Order Com Do sitae — Cine ra ria—Flo we rs

Sleeping — Papilionaceous Blossoms — Clovers —
Effects of Netting—Lamium album—Salvia offici¬
nalis—Fertilisation of Orchis mascula — Bees as
Florists — Conjugation the Door of Progress —
Bees as Fruit-producers — Apples, Raspberries,
Blackberries, Strawberries -— Hints on Culture —
Conclusion.

We are now in a position to examine the modifications
in form and arrangement of the several parts of flowers,
in order to grasp their meaning in relation to fertilisa¬
tion by insect agency. Since some of the most
beautiful aspects of this highly poetical page of the

28 o

BEES AND BEE-KEEPING.

book of Nature depend upon proportion and interfitting
by mutual accommodation, we should be making the
greatest mistake if we confined our attention solely
to the insect which has been the central object of
our investigation ; as well might we endeavour to realise
the beauties of melody by the perpetual sounding of the
key note, or to get harmony and contrast in colour by
banishing all but one. Our bee, nevertheless, will
claim as its right the principal part of our attention,
and it will be its work, rather than the botanical
position of the flower visited, that will determine our
arrangement. Many of the orders, standing widely
separate in systems of classification, have points in
common in relation to their insect fertilisers, and
such may, in consequence, stand side by side.

Very many flowers, in which both anther and pistil
are found, prevent self-fertilisation by maturing one
before the other; and, in the greater number of cases,
the anthers ripen first, such blossoms being called
proterandrous. Since we have already examined the
pelargonium, which is itself proterandrous, let us turn
our attention to the Tropseolum majus (A and B,
Fig. 55), the common garden nasturtium, closely
resembling the pelargonium, and a member of the
same order. Here, as before, the nectar is contained
in a long spur (the nectary, n , A), so long as to
make the flower more useful to humble than to
hive bees. When the flower first opens, the style
is short, and the stigma immature and unreceptive ;
the anthers, also ( a , a, A), are quite unripe, but soon
one or two, as seen in the Figure, begin to rise from
their first position beneath the flower, by an alteration

BEES AS FERTILISERS.

281

in the filament, until they stand just over the stigma,
so that a bee, entering, could not fail to get dusted
on the breast with pollen (now beginning to be shed),
as the tongue is stretched out, and the head pushed
forward to reach the sweet secretion in the spur.
The anthers, continuing to reach maturity, follow their
leaders, one by one, and, during the time that their
pollen is being liberated by dehiscence (gaping of
the pollen pouches), they stand in front of, or close

A, Young Flower (part removed)—a, a. Anthers; s, Stigma; h, Hairs to save
Nectar from Rain ; n, Nectary, or Spur. B, Older Flower—a', Withered
Anthers; s', Receptive Stigma; h’, Hairs; n', Nectary.

to, the stigma. This process occupies from three to
seven days, during which time the flower is, in func¬
tion, male only, although, as carrying both anther and
pistil, it would be classed as hermaphrodite, or of
double gender. The anthers now begin to drop off,
the first to mature being, of course, the first to fade,
and the filaments which bore them, and carried them
into position, now shrivel somewhat, and droop, occupy¬
ing the position shown at a', B. But the style, mean-

282

BEES AND BEE-KEEPING.

while, has grown longer, and the pistil, now adhesive
and receptive, assumes the position, in relation to the
rest of the blossom, which the anthers have successively
occupied. A bee flitting from flower to flower, loading
her legs with pollen, and her honey sac with nectar,
passes, with a well-powdered breast, from the younger
condition (A) to the older (B), and of necessity presses
the pollen grains she carries on to the upstanding
stigma, and cross-fertilisation is accomplished — the
only possible fertilisation, since the two genders do
not co-exist, the blossoms, during their latter period,
being exclusively female.

It is well deserving of notice, that the three lower
petals (one of which has been removed in the Figure)
have their edges cut into a number of narrow strips
0h ), which are turned so as to stand nearly upright.
These refuse contact with water, and perfectly protect
the nectar from dilution by rain, as may be easily
seen by sprinkling water heavily upon one of the
blossoms; but they also appear to serve another
purpose, in compelling the visiting insect to keep its
thorax sufficiently up to bring its hairs on to the
stigma. Looking at the blossom now in the front,
we observe that the lines on the several petals, ac¬
cording to a beautiful and general law in the floral
world, point to the cavity in which the nectar lies,
so that these decorations, enhancing the flower so
much in the estimation of the florist, are, really, so
to speak, guide-posts to the insect visitor.

The sequential movements of the anthers of the
tropseolum are common to many blossoms, and the
explanation now given will make the delphinium

BEES AS FERTILISERS. 283

(larkspur) intelligible, although this belongs to a dis¬
tinct order—the Ranunculaceae. Here, as in the tropae-
olum, the five sepals forming the calyx are brightly
coloured, while the upper one is produced into a long
spur; but, in this case, the two upper petals are con¬
tinued backwards into the spur, and secrete nectar.

The narrow mouth of the flower is surrounded on
all sides by the petals, but these are so shaped, that
the tube they form has an opening beneath, just
behind the entrance. The tongue of the bee, in
stretching towards the nectary, passes, with the head
or thorax, over this opening, into which the anthers,
as they commence to shed pollen, rise, two or three
together, from their position beneath, and so effectu¬
ally powder the insect on the under side. The anthers
drop again when their fertilising dust is exhausted, to
be replaced by others until the last, when the
pistil becomes receptive, and occupies the spot from
which the male organs have retired, thus securing,
as before, cross-fertilisation by pollen from a younger
blossom. To return to our pelargonium (Plate VIII.).
We find this also proterandrous. The anthers ( a , F)
split, and shed their pollen, while the style as yet
presents no stigmatic faces, for the former is now like
a simple rod ; but, when the pollen has wholly or
partly disappeared, the upper end of the style divides
by longitudinal cleavages, and rolls back into view
the five stigmatic, papillose surfaces which had pre¬
viously been mutually protected from possible contact
with pollen. An inspection of a few pelargoniums
in a greenhouse will make these several conditions
absolutely clear.

284

BEES AND BEE-KEEPING.

The order of development noticed in the blossoms
just passed in review is sometimes, though far less
commonly, reversed, as in the Scrophularia nodosa,
or knotted figwort, which, though a most uninviting
plant to the florist, has the charm of solid worth to
the bee-keeper, for, as nectar-producers, its blossoms,
in spite of their ugliness, are hardly to be excelled.
The plant is a strong grower, and loves moist situa¬
tions, where it often attains 6ft. in height, and from
June to October bears, on its square stalks, repeatedly-

A, Young Blossom— s, Stigma. B, Section of Blossom— ca, Calyx; c, Corolla ;
aa, Aborted Anther ; *, Stigma; l. Lip; a, Anthers; n, Nectar; bl, Back Lip.
C, Older Blossom— s, Drooping Stigma; a, Anthers.

forked panicles of flowers. These are somewhat
globular, and very small, not generally exceeding the
size of a pea. Their colour is a dull purplish brown
on the upper petal, passing into a russet green
beneath.

The flower is hermaphrodite, but, as before, the
two genders are . never, actively co-existent. In
this case the stigma is first mature, so that the
name proterogynous (meaning first female) is

BEES AS FERTILISERS. 285

given. When the corolla opens, the stigma ( s, A,
Fig. 56), already adhesive and receptive, presents
itself immediately over the front lip, and bees—
having been dusted by pollen in their visits to older
flowers, and in a manner we shall presently see—as
they reach in after the abundant nectar ( n , B),
transfer this pollen from their hairy breasts to the
sticky stigmatic face. Cross-fertilisation having been
secured, the stigma shrinks and dries, and the style
droops (s, C), while the anthers (a, B), which previously
had been hiding, in a manner which almost looks
like humour, in the pouch-like form given to the front
of the corolla cup for their accommodation in their
moments of bashfulness, now rise into view, take the
place whence the stigma has retired, and begin to
shed their pollen. How singular that the anthers
(a, C) should completely occupy the space over the
lip, arranging themselves in two pairs, so that, in
getting the nectar, the bees must reach across, them, if
the flower is approached in front; while the height of
the back lip ($/, B) is such, that it is impracticable for'
them to steal the honey from behind; and, again, that
the fifth anther {act) is aborted, yielding no pollen,
because it normally stands at the back of the flower,
from which spot the pollen evidently could not be
utilised. As the fertilising dust is carried off for the
benefit of the younger sister blossoms, the yield of
nectar slackens, and the corolla cup at last drops; but
it does not do so until the flower has gratefully given
for others the equivalent of that which it had itself
received. The amount of nectar («) produced is
immense, literally filling the lower part of the corolla,

286

BEES AND BEE-KEEPING.

and often standing much higher than in the illustration.
Some years since, this plant was called in America
Simpson's Honey Plant, and, in a paper extolling its
virtues, a bee was drolly represented as flying aloft,
singing, “ Oh, for a thousand tongues!” Mr. A.
Root says that, watching bees at work upon it, he
saw the nectar actually distilling into a blossom
which, just before, a bee had sucked dry, and that,
in less than a minute, a little bead had been formed.
He states that, as the bees worked, taking up this
thin secretion, they, even whilst humming from flower
to flower, discharged watery fluid ; his opinion being,
that by this process “ they make clear, crystal honey
from the sweetened water, as it were,” that is exuding
so constantly from the nectaries of these little flowers.
This observation of Mr. Root is quite according to
the experience of myself and others. The Malpighian
tubes (page 61), acting as kidneys, excrete rapidly
any excess of water, but the manner in which the
latter passes from the honey sac is not yet clearly
explained.

The honey from scrophularia is only of medium
quality, and it may be urged against the plant that its
appearance is not decorative, and the exposed position
of the nectar, which permits short-tongued insects to
reach it, gives too much encouragement to wasps
and their allies; but all bee-keepers, notwithstanding,
would do well to sprinkle its seeds in waste places.

Nature's fertility of resources is boundless, and the
plan of making hermaphrodite flowers practically
unisexual, by bringing the male and female organs
to maturity at different periods, is often compounded

BEES AS FERTILISERS.

287

with other devices, of which we have an example
in the excellent honey-plant, Epilobium angustifolium
(C, Fig. 57), or rosebay willow herb, belonging to the
order Onagraceas. It is by no means so generally seen
in England as in Scotland, although it abounds in
many parts of Somersetshire. It is common throughout
the cooler parts of the Northern hemisphere, and is
used in Kamtschatka as forming part of a fermented
drink. It grows to the height of 5ft. or 6ft., and
is loaded with racemes, carrying great numbers of
blossoms, which mature in succession during several
weeks. In soils which agree with it, this plant is
even more easily established than removed, as it
creeps along rapidly by lateral shoots. The wild
plants have lilac or pink blossoms, with a lavender-
coloured corolla, and bluish pollen ; but there are three
or four varieties better suited to garden cultivation, of
which, album , roseum and rosmarinifolium are to be
preferred. The last is really a beautiful plant, and
would grace any shrubbery. Some roots were sent me,
through the kindness of Mr. Ingram, well-known for the
interest he takes in bee botany ; they flourished, and,
during fair weather, the flowers were always crowded
with bees. In the Botanical Gardens, Kew, this plant
is grown in the herbaceous grounds, with a multitude
of others, and hive bees are generally found about it
in numbers. It receives its name from the flowers
being placed upon, or at the end of, the pod, which
might almost be taken for the stalk; but its interest
now centres upon its method of fertilisation. When
the flowers open, the style curves backward, carrying
the stigma (j) to the position shown at A, Fig. 57.

288

BEES AND BEE-KEEPING.

The eight anthers now begin to shed pollen, which
bees industriously gather. In two or three days
when the anthers have exhausted themselves, the

Fig. 57.— Epilobium angustifolium (Rosebay Willow Herb), Order
Onagraceoe.

A, Young Flower— s, Stigma turned back; a, Anthers; l, lobe, or Pod.
B, Older Flower— s, Stigma, turned forward; a. Anthers; l, lobe. C, Spike
of Flowers. D, Section of Pollen Grain—e, Extine; i, Intine; ti, Thick
Intine ; /, Fovilla. E, Growing Point of Pollen Grain— e, e, Extine ; i, i, Intine ;
/, Fovilla; pt, Pollen Tube.

style straightens, lengthens to its full dimensions,
and spreads its four stigmas (which previously had

BEES AS FERTILISERS.

289

been shut up together), in the very position to be
pollinated by a bee coming from a younger flower.
The pollen grains (D) demand some little attention.
These are filled with granular particles—the fovilla of
the older botanists—and at the angles the intine is
much thickened, which, as before explained, forms
the covering membrane for the pollen tube (pt, E).

We now turn our attention to a plant yielding a
nectar which has qualities poisonous to human beings,
although it does not seem to be injurious to the bees
themselves. It is a relative of the rhododendrons
and azaleas—of bad repute also, so far as honey is con¬
cerned, although in the same natural order (Ericaceae)
we find our invaluable heathers, whose luscious product
is so highly esteemed. I refer to the beautiful Kalmia
Latifolia (A, Fig. 58), selected both on account of its
peculiar adaptations to insect visits, and because all
apiculturists should know it as a plant to be avoided.
It is a native of North America, growing in damp
places, over very large areas, and is here well known
as a shrubbery plant of great attractiveness, bearing
pink flowers, with the structure of which every lover
of Nature should make himself acquainted. .If a flower-
bud be cut across, the ten anthers will be found to
have their ends tucked into small cavities, or pockets
(ap, D), which appear as bosses on the outside of the
bud, while the filament lies almost in contact with
the corolla; but, as the latter expands (B), the filaments
are bent outwards and backwards, and so brought into
a condition of strain. Now, any sudden jar or rough
handling liberates the anther, when the elasticity of its
filament suddenly throws it up (as at a, C) towards the

2 A

2go

BEES AND BEE-KEEPING.

style, and the pollen ( pg ) will escape by two pores
(po, E), and so- possibly produce self-fertilisation.
But should a bee circle on the wing over the flower,
the legs or under side of the abdomen would first
touch the stigma, and, did the visitor carry pollen,
crossing would be the result. The tongue, now feeling

Fig. 58.- Kalmia latifoua, Order Ericaceae.

A, Flowering Branch. B, Expanded Flower—op, Anther Pocket. C, Section of
Expanded Flower— ap, ap, Anther Pockets; s, Stigma; a, Anther (free); pa.
Pollen Grains m Shower; ca, Calyx. D, Section of Flower Bud—ap, Anther
f Ml t Stamen More Enlar g' ed — 1 a. Anther ; po, Pores ; pg , Pollen Grains ;

its way into the base of the flower, to secure nectar,
would certainly liberate one or more of the anthers,
which, projecting their pollen towards the insect,
would furnish it with material for fertilising some
other blossom. Thus, then, it is that the pollen of
one is carried to the stigma of another. Those un-

BEES AS FERTILISERS.

291

• acquainted with the kalmia would be greatly interested
in liberating the anthers by means of a bristle.

During the celebrated retreat of the Ten Thousand,
as recorded by Xenophon in his “Anabasis,” the
soldiers regaled themselves upon some honey which
they found near Trebizonde, where were many bee¬
hives. Intoxication, with vomiting, was the result.
Some were so overcome, he states, as to be incapable
of standing. Not a soldier died, but very many were
greatly weakened for several days. Tournefort en¬
deavoured to discover whether this account was
corroborated by anything ascertainable in the locality,
and had good reason to be satisfied respecting it.
He concluded that the honey had been gathered from
a shrub growing in the neighbourhood of Trebizonde,
which is there well known as producing the before-
mentioned effects. It is now agreed, that the plants
were species of rhododendrons and azaleas. Lam-
berti confirms Xenophon’s account, by stating that
similar effects are produced by the. honey of Colchis,
where the same shrubs are common. In 1790, even,
fatal cases occurred in America, in consequence of
eating wild honey, which was traced to the Kalmia
latifolia by an inquiry instituted under the direction
of the American Government. Happily, our American
cousins are now never likely to thus suffer, thanks
to drainage, the plough, and the bee-farm.

The beautiful purple heathers of our moorland and
semi-mountain scenery have often given inspiration
to the poet, and lovely indeed are the glowing tints
the countless bells impart to the landscape, as they
reflect the light of the setting sun; but while they

2 A 2

2g2

BEES AND BEE-KEEPING.

thus in mass charm the artist, one can stir emotion
in the breast of the true naturalist. Let us examine
the Erica Tetralix (the cross-leaved heath), which,
though less helpful to the bee than the common ling
{Erica vulgaris , or Calluna ), is very similar to it in
structure. This species, during July and August, in
the southern parts of England, produces abundance
of drooping, wax-like flowers, nearly white at the

Order Ericaceae.

A, Section of Blossom (Magnified Five times)—a, a, Anthers ; ap, ap, Appendages of
Anthers ; /, /, Filaments ; o, Ovary ; s, Stigma ; h, Sticky Hairs. 13, Anther—
f't Filament; ap', Part of Appendage of Anther; p, Pore of Anther, with
Pollen Grains Escaping. C, Blossom (Natural Size). X), Fragment of the
Calyx (Magnified Twenty-five times)— b, b, Simple Hairs ; gh, Glandular Hairs ;
g, g, Glands surrounded by Secretion.

base, and delicately shaded with a rich pink. It is
certainly the most beautiful of our common indi¬
genous heaths, and grows freely on moist, mossy
ground and bogland throughout the kingdom. Open¬
ing the bell, which sways in the wind mouth down¬
wards, we find a straight, pinkish style, terminated
by the stigma (j, A, Fig. 59), hanging in the centre

BEES AS FERTILISERS.

293

like a clapper, the stigmatic face in large part closing
the narrow opening. The filaments (/, f), eight in
number (all but two are removed in the Figure, to
avoid confusion), start from the base of the ovary ( 0 ),
where the nectar is secreted. These filaments, like
those of kalmia, act as springs, but, in this case,
their function is to hold the anther close against
the rod-like style. The anthers, also, are provided
with oval openings, or pores ( p , B), which are placed
at their lower ends; but since they are held side by
side, in a circle around the style, the pore of one is
opposite to the pore of the next, so that the escape
of pollen is prevented. Each anther consists of two
cells, and each of these is furnished with a horn-like
process ( ap, A), expressly intended to be in the way
of the tongue of the nectar-gatherer. She arrives,
but the opening is too small to admit her head, and
the distance from the mouth of the bell to the sweets
sought is as far as she can reach, so her head is
brought up into contact with the viscid stigma.
Here she leaves her load of pollen (for she wears
hair powder, as we shall see, while she is at work
on heather), and so accomplishes her work as fertiliser.
The tongue, as it runs up, must strike one or more
of the sixteen anther appendages, which act like
levers, and so disarrange some of the anthers them¬
selves, and separate their pores, when down rains
the fertilising dust upon the bee’s little brow (where it
remains, as it is beyond the reach of her leg-brushes,
so that she gathers no pollen from heather). She
sucks her nectar, and passes to the next blossom ; the
elastic filaments restoring the anthers to order, and

294

BEES AND BEE-KEEPING.

awaiting another visit; and the bee. applies, almost
as quickly, the pollen she carries, to the receptive
stigma of the next bell. How clear it is, that some
correlation in size between bee and flower is needed !
Tiny insects might creep into the bell, and, passing
up its sides, secure its nectar, without touching the
anther appendages; but if they did, they would not
be dusted, the pollen would be but wasted, and, worse,
the nectar would be gone, and the bee would not be
encouraged. A singular device prevents this : the
plant, especially about the flower stalks, bracts, and
calyx, is covered by glandular hairs, or trichomes ( h ,
A, and gh, D), which constantly yield a viscid body,
that would stick fast the little thieves if they ventured
to attempt to make themselves guests without an in¬
vitation. These clammy hairs, however, trouble not
the bee : she grasps the glossy bell with her legs,
and does not touch the calyx. Her weight but
makes the position of the flower the better for the
proper placing of the pollen. In the calluna (the
ling), the flowers are more horizontal, and the style
curves upwards, so that the bee’s tongue is inserted
beneath it; but the whole plan of action is similar
to that occurring in Erica Tetralix or E. cmerea. The
sticky hairs, however, are not present, while both the
latter species possess them—and why? The mouth of
the blossom is so small that it is its own protection.
The window need not be shut if its opening is
covered by narrow bars, and Nature does not waste
force in defending that which needs no defender.

I was greatly pleased, some time since, in studying
the blossom of the Arbutus Unedo (strawberry tree),

BEES AS FERTILISERS.

295

and, although the flower is not especially interesting
to bee-keepers, I venture to give ray results, as illus¬
trating and explaining much that has already been
said. This tree-like shrub belongs to the same order
as the foregoing, and is characterised by bearing a
berry containing many seeds ; and its resemblance to
the well-known fruit gives the popular name. It is
a native of Southern Europe, but with us it is a hardy

Fig. 60.—Flower and Details of Arbutus TJnedo (Strawberry Tree),
Order Ericaceae.

A, Section through Blossom (Magnified Five times)—/, /, Filaments of Anthers ;
a. Anther driven from Style; a', Anther in Natural Position; ap, Appendage
of Anther; s, Stigma; 0, Ovary. B, Blossoms (Natural Size), s', The Five
Stigmatic Faces communicating with the Five-celled Ovary, more Magnified.

evergreen, reaching 16ft. high, or even more. Its
blossoms are greenish-yellow, and are formed in
October and November, and, with the same general
plan of structure as those just noticed, it has some most
singular differences and modifications. We find here
ten anthers^ placed around a straight style, by spring¬
like filaments ; but the former are disposed in two rows
of five each. The inner has its pores placed on the

296

BEES AND BEE-KEEPING.

end, instead of the side, of the cell, so that the style
itself stops the orifices ; upon the backs of the inner
anthers the outer row is pressed, and thus a ring of
twenty appendages ( ap , A, Fig. 60) is set round the
centre of the flower. The stigma is receptive before the
anthers are ripe. Unless insects quickly visit the
blossom, the outer anthers drop back, as at a, A; but
all the pollen does not at once run out, for the anther
is lobed within, and is delicately poised at the back,
by a very slender termination to the filament. The
pollen which falls is here held by the inner hairs of the
corolla (shown in the Figure), and may be swept up by
bees' tongues. The second set of anthers (a'), at a
iater period, fall back, and sprinkle their pollen as
the others. The suggestion seems to be, that, if
bees do. not come, then smaller insects, walking up
the corolla inside, may get coated, and so fertilise
other blossoms (for we must remember that these
flowers are proterandrous) by walking down their
styles. For it is singular that, notwithstanding the
large mouths of the corolla, no sticky hairs occur;
but their office is, in this case, performed by the
filaments (/), which are enormously thickened, and
covered with thin, cottony hairs, so that the ten so
fill the upper part of the bell, that very small insects
could not force their way through them to get the
nectar.

It is in this useful order that we find also the
cranberry, bilberry, and whortleberry, all the blos¬
soms of which could be well made out by applying
the above explanations as an examination might
warrant.

BEES AS FERTILISERS.

297

The consideration of the movements of the fila¬
ments of Kalmia and Arbutus Unedo naturally intro
duces a common British plant, whose filaments are
remarkably irritable, and hence often secure cross¬
ing. In the common barberry, which, in June,
bears drooping racemes of yellow flowers, the six
filaments spread directly outwards, standing just over
the six petals, which bear twelve conspicuous honey
glands that are very alluring to bees. Should one
of the latter, in seeking sweets, touch a filament,
it immediately springs upwards, striking the insect, so
as not only to dust its body, but to so startle it that
it retires to another flower, when the pollen carried
off is immediately transferred to the receptive edge
of the upstanding stigma. So persistent is this curious
property, that the filament will contract upon being
touched after its removal from the flower.

We have previously noticed several cases in which
the genders appear on different ( dioecious ) plants,
the flowers being unisexual, and, in consequence,
incapable of self-fertilisation; and we have now to
consider a most interesting set of variations, in which
the flow'ers become practically dioecious, although
they remain hermaphrodite, securing cross-fertilisa¬
tion by differentiating into two, or even three, dis¬
tinct forms, which are complementary to one another.
If, by example, a handful of primroses be gathered
promiscuously from several plants, they will be found,
upon examination, to present very apparent dissimi¬
larities amongst themselves, some having a pale green,
almost globular form (s, A, Fig. 61), the stigma, at
the top of the corolla tube, others, at the same spot,

BEES AND BEE-KEEPING.

showing five anthers (a', B), nearly closing the mouth.
Upon splitting these two forms, the first will present
an enlargement in the centre of the corolla tube,
and here the anthers (a, A) take their position. The
second has an enlargement at the top, where the
anthers ( a ', B) are placed, while the centre of the
tube is destitute of any widening, but contains the
greenish stigma (s'). It is clear that, if a bee,
probably a Humble, visits the first blossom (A), the

A, Long-styled Flower— s, Stigma ; st, Style; a, Anther; o, Ovary; pg, Pollen
Grains, more Magnified. B, Short-styled Form— a', Anther; s’, Stigma ;
sf, Style; o', Ovary ; pg', Pollen Grains, more Magnified.

long tongue, fully outstretched to get at the nectar,
will be coated with pollen upon the centre of its
length ; and should the bee now pass to the second
form, the carried pollen will be in the correct posi¬
tion for fertilising the flower, while the tongue will
get coated at the root, for subsequently pollinating
the first form. Darwin, in a series of admirable ex¬
periments, proved that, although seed might be pro¬
duced artificially in the plant under consideration, by

BEES AS FERTILISERS.

299

putting pollen from form A upon the same or another
flower of form A, that the best seed, and the largest
number per capsule, could only be obtained by cross¬
ing, not only two flowers, but also the two forms,
which are naturally bound together by mutual depend¬
ence. This leads to the inquiry, Does any ascer¬
tainable difference exist between these pollens ? Micro¬
scopic measurements show that the pollen grains of
form A (pg) only contain about one-third the material
of those of B, seeming to indicate, although Darwin
hazards no opinion here, that the larger grain is best
suited to forming the longer pollen tube required in
the long-styled form. By further experiment, the
existence of that which is commonly called prepotency
was proved— i.e., pollen placed on the stigma of the
flower form whence it had been derived, would be
rendered powerless by subsequently adding pollen from
the complementary blossom.

Dimorphism (or double form) is more common than
even botanists, not long since, suspected, and amongst
dimorphic plants we find those of the highest utility to
the bee, because to many such the bee, or some other
insect, is a sine qua non. In the Linacese , or flax
family, e.g., certain species are only capable of pro¬
ducing seed at all when intercrossed. In Linum grandi-
florum (A, Fig. 62), the stigmas (s, s ) and anthers (a, a )
are so placed, that intercrossing must be generally
brought about by bees reaching after the nectar
secreted, at five points, at the outside of the anther
bases; and experiment has fully shown, that if the pollen
of B be placed on its stigmas, or those of any other
similarly formed flower, not only is fertilisation not

3 °°

BEES AND BEE-KEEPING.

effected, but the pollen utterly fails in even forming
a pollen tube. The pretty Pulmonaria officinalis , or
lungwort, presents another example of the same
singular fact, which obtains, in a less degree, with many
other plants ; Darwin, in one list, enumerating thirty-
four. Polygonum fagopyrum (buckwheat), previously
figured, is strongly dimorphic, having short styles and
long filaments, or the converse.

But plants occasionally, as previously hinted, present
three forms in the same species. No example is more
surprising than that of purple loosestrife (.Lythrum
Salicaria ), which, withal, is a good honey-producer,
secreting nectar all round the base of the ovaries, and,
in consequence, visited with great frequency by hive
bees. It is also amongst the most handsome of our

BEES AS FERTILISERS.

3 0:

native perennials, with its long, tapering spikes of crim¬
son and purple, borne on steins 3ft. or 4ft. high, and
decorating gaily the banks of the stream, where it holds
its own amongst the sedges, rushes, and sallows.
Although a water lover, it may be naturalised in the
garden ; and I know of no plant likely to afford so
much pleasure to the scientific bee-keeper. It has,
in all its forms, twelve anthers, arranged in two rows
of six, and a centrally placed pistil. But the lengths
of the style and filaments in no two forms agree; when
the style is short, the stamens are medium and long;
when the style is medium, the stamens are short and
long; and when the style is long, the stamens are short
and medium. The long pistil is fertilised by the long
stamens of the other two forms, the medium by the
medium, and the short by the short; for, as bees pass
from plant to plant (each plant bearing only one form
of flower), the pollen finds its proper resting-place by
the position given to it by the anther whence it
came, for the long, medium, and short stamens touch
the bee on different parts of the body, which are
subsequently applied to the long, medium, and short-
styled stigma respectively. The wonders of this
interfitting are too many for description. Even the
pollens are truly diverse, and of three kinds, while
the filaments are distinct, the long being deeply red,
the medium and short white. The anther cells of the
long filaments are nearly black, and the pollen grains
a brilliant emerald green, while the pollen of the short
and medium anthers is yellow. The green pollen
grain is large, the yellow pollen of the medium anther
smaller, and that of the short anther less again, seem-

302

BEES AND BEE-KEEPING.

ing to indicate, as before, that the pollen grain has
its size accommodated to the length of tube it has
to produce. The bee-keeper possessing these plants
will never lack a source of amusement and in¬
struction both for himself and his friends.

The order Compositae , which embraces no less
than about 10,000 species, of which 113 are British,
includes very many plants that are of the highest
utility to the bee-farmer. The name of the order
implies that the flower-head ( capitulum ) really carries
many blossoms, which, on account of the closeness of
their packing, would popularly be regarded as one.

Fig. 63.— Section of Flower-head (Capitulum) of Common Cineraria, Order
Compositce (Magnified Four times).

A, Ray-floret; B, Receptacle, with Involucre—1,1', 2, 2', &c., Florets Entire and
in Section, in similar Conditions of Development.

Let us first examine a capitulum—and, for our pur¬
pose, a sunflower, a field daisy, a thistle, or a
golden rod, with its rich nectar, would have answered
perfectly; but I choose a cineraria from the green¬
house, for the same reason that I previously selected
the pelargonium. Looking at the flower-head, we
see a ring of petals surrounding a convex centre,

BEES AS FERTILISERS.

303

which careful inspection will show to be formed by
the upper ends of a number of tubular flowers.
Making a cross section, we find the central flowers,
or florets, less developed than those at the side ;
so that, by passing from the centre to the cir¬
cumference, we trace the steps of progress through
which each floret must pass. In No. 1, Fig. 63, the
extension of the corolla tube at its lower end, to ac¬
commodate the ovary, is seen, while above, the corolla
is just opening. About a day later this floret will
have assumed the form of No. 2, where the anthers
have grown up partly into view. In this order the
anthers and their filaments surround the style, the
anthers uniting at their edges, so as to form a tubular
sheath, into which the pollen is shed. The style fills
the tubular sheath in its lower part, like the rod of a
popgun, and, as it grows, drives the pollen before it,
until at last the anther tube gives way, and the
pollen is pushed out as we see it at Nos. 4, 4', Figs.
63 and 64, when insects, searching for sweets, and
collecting or eating the pollen, will get dusted
beneath. The style still grows on, and now appears
(5) having its end covered with filamentous hairs,
which have acted the part of a chimney sweeper's
machine in driving the pollen before it. A day or
two later, when its fertilising dust has all gone, the
style splits, and curves back (6), so as to expose
the now receptive stigmatic faces; and thus, as in so
many similar cases, crossing is. secured. Coming
now to the outside floret, carrying one of the rays
which make the external ring of the flower-head, we
find a distinct alteration. Here the style carries no

304

BEES AND BEE-KEEPING.

brushes, for this floret is purely female, having i
anthers. No pollen requires sweeping out, and, thei
fore, brushes, which could effect no purpose, a
not formed ; and again, had this floret produced polle
it could not have been utilised, at least on the san
flower-head, since all the florets are proterandroi
the younger in this matter serving the elder, and
absolving the eldest of all from the need of polle

OF fart of One of the COMPOSITE (Cineraria), Magnified Ten times;
Lettering as Fig. 63.

production. Although the plan throughout the order
is so similar that the comprehension of one example
gives the key to every other, yet there are variations
of detail which must be remembered. Thus, in some
composite flower-heads every floret bears a ray, as in
the dandelion; or all may be tubular and perfect, as
in the spear thistle; or the outermost florets neuter,
as in the corn bluebottle; or female, as in the case just
examined; or monoecious, those of the disk beinp- male.

SLEEPING FLOWERS.

305

and the rays female, as in the marigold; or even
dioecious, one plant bearing male heads, and another
female, as in the cudweed ( Gnaphalium dioicum ).

But all are wonderful; the commonest of composites,
the field daisy, peeping out amidst the grass though
it forms but little dots in the pattern of Nature’s soft
carpet, is still a world of wonders in every one of the
hundred tiny tubes that make up its little face, with
its frill of white and pink. It carries within itself
the counterpart of all hitherto explained, with marvel
upon marvel beside, to reward him who seeks; for
Nature is never truly wooed that she does not lift
her veil and smile. The daisy, too, is a sleeper;
when the sun goes down, she closes her ray-florets,
to open them again when day returns; and hence
her name —“ day’s eye.” So with other flowers—
the dandelion, the hawkweed, the sandwort, the pim¬
pernel, all of which choose special hours to sleep
and wake, adding a day nap when clouds hide
the sun. Some blossoms are like Convolvulus sepium
(the great bindweed), which closes at night, unless
the moon is shining, and the stillness is broken by
the low murmur of the moth’s wing, when it remains
open. If to this we add, that wind-fertilised blossoms
do not sleep, and that those depending on moths are
especially fragrant during the most active hours of
these nocturnal insects, is not the conclusion almost
irresistible, that these peculiarities have relation to the
modes of fertilisation of each ?

No order contains a larger proportion of plants of
utility to bees than the Leguminosse, every British
representative of which has an irregular flower, of

2 B

3°6

BEES AND BEE-KEEPING.

papilionaceous (butterfly) form (A, Fig. 65). It is
probable, that all flowers having an irregular corolla
are adapted to fertilisation by insects, and that the
latter are prevented, by the irregularity, from reaching
the nectary, except from that position which makes
their visits effective in securing a cross. In the
papilionaceous tribe, it frequently happens, that the
nectar-gatherer, in alighting, causes certain mechanical
changes by its weight, by means of which pollen is
transferred to its body for distribution to neighbouring
blossoms. In this order we find lucerne, sainfoin,
melilot, clover, vetches, and many others ; but the
Pisum sativum, (the kitchen pea), an importation from
Southern Europe, is an excellent typical flower, although,
singularly, in our own country, failing its native insect
attendants, it has acquired the power of self-fertilisa¬
tion.* The corolla has five petals—a large upper one
( v ), the vexillum, or standard; two that are lateral,
the alae ( al ), or wings ; and two, more or less united
at their lower margins, forming the keel, or carina,
which, within its boat-shaped cavity incloses the
stamens and pistil. The anthers are ten in number; the
filaments of nine of them (a', C) are confluent, form¬
ing a covering for the ovary beneath and at the sides,
but slit above, where the tenth anther (a), with its
isolated filament, is placed. This slitting gives the bee’s
tongue access to the nectar at n, B and C. The style is
somewhat hairy, while the anthers open early, and dis¬
charge their pollen, which mainly lodges upon the
style. If the blossom of a pea, or vetch, be taken

* Rev. G. Henslow on “ Self-fertilisation of Plants ” (Transactions of
Linnean Society, 2nd Series, vol. i., page 361).

BEES AS FERTILISERS.

307

to pieces, it will be seen that the keel petals have
each a protuberance, which fits into a corresponding
hollow on the inner sides of the alae, so that the
latter cannot be depressed without carrying the keel
with them. This is really what happens when the
bee settles. The style, forming part of the rigidly
set pistil, relatively rises between the keel petals,
and touches the bee on the thorax, as at B, leaving

Fig. 65.—Papilionaceous Blossoms, and their Method of Fertilisation.
A, Expanded Pea Blossom, Order Leguminosas—v , Vexillum ; al, Alae with Carina
between. B. Partial Section of Flower of Vetch being Fertilised by Cyprian
Bee (Magnified Twice), Right Ala removed below line a, 6—1/, Vexillum;
n', Nectar Gland ; al' Ala ; c, Carina containing the Pistil, the Stigma of which
is striking the Bee’s Breast. C, Section of Pistil, showing Ovules (Peas) in
Ovary— n, Nectary; a, a', Anthers ; s, Stigma.

some pollen, which may be placed on the stigma of
the next flower. All this can be easily seen, by hold¬
ing the blossom steadily, and pushing down the alae
with the fingers. At the departure of the insect, the
style again retires, to repeat the process if necessary.
The stigma first touches the bee’s body, so that crossing
is brought about; and then, as the tongue is employed
in sweeping up the nectar, a new supply of pollen
is given ; so that, in visiting a succession of blossoms,
the pollen of one is transferred to the stigma of the
next. The description now given applies to Lathyrus

2 B 2

3°8

BEES AND BEE-KEEPING.

and vetches, but in the scarlet runner a singular and
interesting modification must be noticed. The keel
is prolonged into a narrow snout, which is literally
coiled like a snail’s shell, and through which the style,
similarly twisted, is prolonged. When a bee visits,
the stigma first makes its appearance, and then the
pollen-coated style, acting in the manner previously
noticed. The French bean has a structure identical
with that of the scarlet runner, and both yield honey;
yet the latter is sterile without insect action, while
the former, where no insect can fertilise it, may be
forced, yielding seed in full abundance by self¬
impregnation. In the melilots and trifoliums, and
Onobrychis (sainfoin), we have the plants which most
gladden the bee-keeper’s heart. Their honey is of
rare quality, and its amount is astonishing. Their
fertilisation is accomplished as before, with slight
differences, since the style is not brush-like, but the
anthers themselves pass out of the keel to give up
their pollen. Darwin points out the utility of bees
to these plants. The flowers of Trifolium incarnatum
(crimson trefoil), which were visited by bees, pro¬
duced between five and six times as many seeds as
those that were protected (covered with a net). Of
Trifolium pratense (common purple clover), ioo
flower-heads on plants protected did not produce a
single seed, whilst ioo heads, on plants growing out¬
side, which were visited by bees, yielded 2,720 seeds.
In Trifolium repens (white Dutch clover), the crossed
and self-fertilised plants yielded seeds in the ratio of
ten to one ; and, in another experiment, twenty heads,
unprotected, yielded 2,290 seeds, while twenty pro-

BEES AS FERTILISERS.

309

tected heads had “only a single aborted seed.” The
Trifolium repens , taking first rank as a fodder plant, as
well as a honey-producer, delights in chalky ground, and
often the powdering of lime on the soil will cause a
crop of it to appear where previously it had not
been cultivated, or known to exist.

The sweet-scented Labiatae are not unworthy com¬
panions of the Leguminosae , for not a single plant

d d'

A, Side View of Flower—c, Calyx; l, Labium, or Lip ; s, Stigma ; b, Narrow Mouth
to Corolla Tube ; d, Upper Lip arranged as a Hood. B, Front View of Part of
Blossom— d'. Upper Lip ; a. Anthers arranged in Line; s', Stigma. C, Side
View of Stigmatic Faces (s") carrying Pollen Grains. D, Enlarged View of
Anther dehiscing— pg, Pollen Grains; h, Hairs holding Pollen Grains.

of the order possesses poisonous properties, whilst
amongst them we find lavender, thyme, rosemary,
the mints in their varieties, marjoram, sages, sweet
basil, savory, balm, germander, horehound, &c., most
of which give special aromas to honeys gathered
where they abound. Lamium album (the white
dead nettle), to be found in every hedgerow in the
South of England, will make evident the general

3io

BEES AND BEE-KEEPING.

characters of the order; but the size of the blossom
fits it for fertilisation by Humble bees. The extra
size, however, will aid us if we pull a flower to pieces.
The nectar is found in the lower portion of the tube,
around the ovary, where it can easily be seen by
removing the calyx ( c , A, Fig. 66), and slitting the
corolla. This is protected from rain by the umbrella
form of the upper lip, which acts more perfectly be¬
cause it is surrounded by hairs. The tube-like portion
of the corolla throws out a broad lip (/), which serves
as an alighting-place. The flower, in its earlier stage,
has the four anthers (for one is aborted, as in scrophu-
laria, and for the same reason—see page 285) arranged
in line just under the hood, or umbrella (a, B), so that,
as the bee stretches in after the honey, the central
parts of the head and thorax get coated with pollen.
We notice another curious adaptation : the anthers are
very hairy (D), while they look towards one another,
so that, as the fertilising granules leave the anther
cells, they are held by the hairs right in the median
line. The pollen is now carried away by the bee,
to be transferred, in turn, to a more advanced flower,
where the stigma (s", C) is both receptive and pro¬
minent. Within the corolla, towards the base, we
find a ring of hairs pointing upwards, which effectually
prevent small insects, whose backs could not be applied
to the anthers, from creeping down the tube, and so
stealing the nectar.

In several other species of Labiates we have still
more singular modifications, cross-fertilisation being
secured, as in many previous cases, by the stamens
coming to maturity, and shrivelling before the

BEES AS FERTILISERS.

stigma is receptive; and, in addition, mechanical
means are used for placing the pollen on that part
of the fertilising insect which alone can be effective.
If we take a recently-opened flower of the Salvia
officinalis (common sage), A, Fig. 67, and make a sec¬
tion, we shall find the stamens are of a most modi¬
fied character ; the filament (f C) is extremely short,
and very stiffly set upon a curiously modelled part
of the corolla, apparently specially designed to give
rigidity (s, E). The anther cells, instead of standing
at the end of the filament, are widely separated, by a
long white rod (c, C), the connective, which is itself
hinged to the top of the filament, while the lower
anther cell is aborted, producing no pollen. The
nectar is secreted, as in the lamium, at the end of
the corolla tube. When the blossom is entire, the
two aborted anther cells meet together, and bar the
entrance to the flower, at the lower part. A bee,
attempting to enter, drives her head against the
aborted cells, which immediately yield, and run back
into the flower, turning the connectives on each side
on their hinges {hi, D), at the end of the filaments, by
which the anther cells ( a , C), carrying pollen, are patted
down on to the bee's back ( a , D); and here a dense
patch of coloured dust is left. The nectar having been
absorbed, the bee departs, and, as its head is withdrawn,
the connectives revolve into their old position, and the
anthers await another arrival, until all the pollen has
gone. While the anthers wither, the style {si> C and D),
short in the young stage of the flower, grows rapidly,
and presently occupies the position seen at B. Our
gatherer arrives, decorated on the back, and, pushing

312

BEES AND BEE-KEEPING.

into the corolla mouth, gives up the burden, of
which it is unconscious, on to the stigmatic face,
now waiting to receive it. If a pencil be passed
into the mouth of a flower of Salvia patens or
S. fulgens —both of which, on account of their great
length of corolla, are desirable for illustration—the

Fig. 67.—Blossom of Salvia officinalis, Order Labiates (Natural Size).

A, Young Flower, showing aborted Anther Cell. B, Older Flower, showing Stigma.

C, Section of Young Flower—a, Anther Cell; ac, Aborted Cell; c, Connective ;
/, Filament; hi, Hinge of Filament; co, Corolla ; ca, Calyx ; st, Style ; s, Stiff
Attachment of Filament l, Labium ; A, Interior Hairs ; nr/, Nectar Gland.

D, Section of Young Flower, with Bee entering; Lettering as before. E, Section
of Base of Flower flattened out, Lower Part shown—st, Style ; s and /, Stiff
Attachment of Filament of Pollen-bearing Anther ; aa, Aborted Anther;
A, Interior Hairs ; ca, Calyx ; ng, Nectar Gland.

descent of the anther, attached to a connective,
perhaps an inch in length, and looking like a mimic
chopper, the blow of which decorates the pencil,
cannot fail to cause astonishment. Indeed, I can
hardly imagine any sight more curious than that pre-

BEES AS FERTILISERS.

313

sented, any summer afternoon, in the Herbaceous
Garden at Kew, where hundreds of various Labiates,
growing side by side, attract crowds of bees, of
different genera and varied sizes, which, entering
the curious flowers these plants bear so profusely,
bring the hiding anthers into view, literally getting
patted on the back for their pains. It is only,
however, insects of the correct bulk that can effect
the work; they must be large enough to meet the
anther, and reach up to the stigma; and so, to prevent
smaller ones from stealing that abundant nectar in¬
tended for their superiors, in size at least, the lobster-
pot arrangement, previously noted, is supplied, and
seen at h, C and E, up to which the nectar frequently
extends. But this is not the only manner in which
thieves are kept at bay ; in a salvia common on the
Continent (Salvia glutinosa ), there are no internal
hairs, but the flower-spikes, bracts, and the entire out¬
side of the blossom, are covered by secreting trichomes,
which are exceedingly sticky, frequently holding as
prisoners crowds of small insects, that vainly paw
the air, pleading for release. The inside of the
flower, including the lip upon which the bees settle,
is not adhesive, so that the favoured insect is not
incommoded. The plan brings to mind the Erica
Tetralix (page 294).

In the order Cruciferae we have many useful
honey plants, embracing the wallflower, stock, cress,
rocket, cabbage, turnip, and mustard ; the wild form
of the latter (charlock, or cadlock), as a widespread
weed, yielding, in some districts, the staple of the
bee-keeper’s harvest. The most usual form of adap-

3 I 4

BEES AND BEE-KEEPING.

tation in this order, which is very varied, is one of
great interest, and must be ascertained by an in¬
spection of the flowers. The anthers in the young
blossoms face the style, but before they ripen they
turn their backs, and shed the pollen, which is thus
in the least likely position to find its way to the
stigma of the flower yielding it, but in the most
favourable place for adhering to insect visitors acting
as cross-fertilisers. The retrorse anther, as it is
called, is frequent in its occurrence in other orders.

Our space has permitted us to deal with types only,
and these would be extremely incomplete without
some notice of orchids, which have always been
objects of wonder, but have never attracted more
attention than in recent years, as their investigation
has revealed devices which appear to the last degree
romantic. The one example chosen for illustration
is a British species, Orchis Morio , which, as I have
several times witnessed, is habitually visited by the
hive bee, and so here is of deeper interest than the
more extraordinary exotics, many of which are large
nectar-producers, while all species of the sixteen
British genera are of only moderate value in this
respect. The flowers in this order are exceedingly
unlike those we have previously studied, so that
some little attention must first be given to general
structure. In Fig. 68, A, we find, as in all common
orchids, but one developed anther (a), which has no
distinct filament, for this is confluent with the pistils,
forming together the column—the part of the flower
immediately in front of the bee’s head. The anther,
which we have seen in other cases to carry the

BEES AS FERTILISERS.

vivifying element (the pollen), in the form of granules,
collected in two cells, has here a peculiar structure :
its cells are two, but they are so widely separated
(a', a', C) as almost to appear like two separate anthers ;
while the pollen they contain coheres in masses
(pollinia, po , D), held together by internal elastic

Fig. 68.—Orchid (Order Orchidacece) Blossoms and Details.

A, Flower of Orchis Morio, Sepals, two Petals, and side of Spur removed, with Apis
Mellifica (op), Hive Bee, sucking Nectar— a, Anther; po, Pollinium or Pollen
Mass ; r, Rostellum ; si, Stigma (side view); l, Labellum ; os, Ovary ; n, Nectary ;
hr. Bract. B, Bee fertilising Orchis Morio—a. Anther with Pollinium removed ;
po, Pollinium, attached to Bee’s Head and applied to Stigma ; other Letterings
as before. C, Front View of Orchis Morio, Magnified Three times. Sepals and two
Petals removed— Ir, Lip of Rostellum ; f, f, Fissures in Front of Anther Cells
(a', a '); other Letterings as before. D, Pollen Masses, &c.—po, Pollinia ; c, Cau-
dicle ; vd, Viscid Disc ; vg, Viscid Globe ; Ir, Lip of (r) Rostellum. E, Head of
Bee, carrying (po ) Pollinium— an, Antenna;. F, Position of—po, Pollinia (thirty
seconds later), partially depressed. G, Head of Bee—an, Antenna" ; po, Pollinia
(sixty seconds later) fully depressed. H, Pollen Granules (much magnified),
held in packets by thin elastic threads. I, Head of Bee, carrying (po) Pollinia
of one of the Vandeoe — an, Antennae.

316

BEES AND BEE-KEEPING.

threads, which also tie each mass, at the end of a very
curious, stalk-like appendage (the caudicle, c), which
is again attached to a viscid piece of membrane (the
viscid disc, vd), having below it the viscid globe
{vg). The method of distributing the pollen will come
before us presently. The stigmatic faces are, theo¬
retically, three, but only two are fertilised by pollen,
with the formation of pollen tubes, entering the ovary
in the usual manner; their sticky faces are seen, side
view, at si, A; in the front view, si, C. The third
stigma is modified into what is called the rostellum
( r , A), which contains the viscid matter of the discs
and globes just mentioned, playing a most whimsical
function, in order to secure crossing. The outer
portions of the flower consist, as in most orders,
of calyx and corolla, here divided into three sepals
and three petals respectively. All of the former,
and two of the latter, have been removed, to permit
of an uninterrupted view of the organs of reproduc¬
tion. The third petal, properly the upper one, but
made the lower by a semi-twist of the ovary (ov, A),
is larger than the others, and offers a landing-place
to insects, as we see by the position the bee has
taken. It is called the lower lip, or labellum (/), and
is carried backwards in the form of a spur, where it
assumes the functions of a nectary * (n), and so
attracts visitors. The anther cells are longitudinally
open in front, the fissures (/, /, C) of the covering
occurring before the flower opens, so that the pollinia
may be taken out from their pouches, in which they
lie, but to which they are not attached. The
membrane, forming the whole external surface of the

BEES AS FERTILISERS.

317

rostellum, is at first continuous, but, as soon as the
flower opens, the slightest touch causes it to rupture
transversely, in a sinuous line ( Ir , C and D) in front
of the anther cells. Let us now suppose that a bee
(ap, A) alights on the labellum, and advances the
head, in order that the tongue may reach the nectary
(#), where the sweet liquid must be secured by an
abrasion of the delicate lining membrane. The
rostellum, irritated by the touch of the insect, im¬
mediately ruptures its covering skin (if this were
entire at the previous moment), and the pushing
forward of the head depresses its lip ( lr , D), so
that the viscid discs (vd), formerly a part of the
covering membrane of the rostellum, and the viscid
globes ( vg ), are exposed, the latter infallibly coming
into contact with some part of the bee’s head. So
viscid are these globes that they firmly stick to what¬
ever they touch; moreover, they have the property
of setting hard in a few seconds. During the time
occupied in sucking the nectar, they, in consequence,
become firmly attached to the head of the bee, the
connected pollen masses still lying in the anther
pouches, whence, as we know, they can be readily
withdrawn ; this is accomplished as our bee retires
carrying a decoration in the form of two upstanding
yellowish-gredn horns ( po , D and E, the pollinia of
the orchid). Darwin pointed out that all this may be
exactly imitated by a well-pointed pencil, or a stiff
bristle; and none would regret the little trouble in¬
volved in growing a few common hardy orchids, in
order to have the pleasure of showing the experiment
to friends. But how are these pollinia to be made

3 i8

BEES AND BEE-KEEPING.

effective ? How is their material transferred to the
surfaces of the stigmas; for will not the next flower
visited have these masses thrust forward towards
its own anther pouches ? Such would be the case
with the visits immediately succeeding the first.
Watching our pencil point, or the head of the bee, the
pollinia, at first erect on their caudicles {po, E), and
firmly secured by the drying of the viscid globes, begin
to incline forwards, and continue to move, always in
one direction (towards the pencil point), until they have
swept through an arc of about 90°, finally standing
as at po , G; the movement occupying about thirty
seconds on an average. Re-inserting our pencil,
or our bee making another visit, will now secure
fertilisation, because the pollinia immediately strike
the stigmatic faces {st, C), as we see actually
being done at po, B. How perfect the adaptation !
But where lies the secret of the movement executed?
The little viscid disc {yd, D) is endued with a power of
unequal contraction, and produces the required change
in position, the time occupied by it permitting the bee
to get from one plant to another, so that the best form
of crossing is secured. And yet another adaptation
demands attention. The pollinium is very coherent;
but the elastic threads holding it together in packets
(H) break with the energy, the insect can exert, so
that some pollen is left, and yet a mass carried away,
which may be effectively used upon flower after flower,
until at last the ragged caudicle alone remains.

Some time since, when I had announced the discovery
of some diseases previously not known amongst bees,
a bee was sent to me, whose portrait is given at I,

BEES AS FERTILISERS. 319

and I was asked to name the disease which had caused
an abnormal outgrowth upon its head. It had visited,
there can be little doubt, an orchid-house, and had
carried away a decoration from one of the Vandeae.

The description given above applies to Orchis
mascula, fusca, maculata, and latifolia, as well as
Aceras anthropophora (the man orchis), in all of which
the pollinia undergo the same curious movements of
depression which are necessary to enable them to
strike the stigmatic surfaces. The behaviour of these
British specimens is quite commonplace, however, in
comparison with that of some exotic orchids explained
by the genius of Darwin. A few words in reference
to one will suffice. In Catasetum, the genders are
divided, and the male blossom is provided with a
strange pair of long additions, called by Darwin “ the
antennse.” In some species both are equally active,
but in others the right one seems functionless, while
the left is intensely irritable, and, should an insect
touch it, a vibration is transmitted to a certain mem¬
brane, which is instantly ruptured, and so sets free
the pollen mass, which is shot forth from the extremity
of a liberated spring, viscid end first, to attach itself
to the back of the insect. The startled bee flies,
possibly, to the female flower, and here accomplishes
a cross ; this act being favoured by the curious habit
of the Apidae of visiting one kind of flower only,
during any single excursion—a habit for which no
sufficient explanation can be given, although it will
receive further notice hereafter. Sir John Lubbock says:
“On one occasion, Darwin touched a male catasetum
in my presence, when the pollinium was thrown nearly

320

BEES AND BEE-KEEPING.

3ft., and stuck on the pane of a window.” I have
seen this droll performance at Kew, and although 3ft.
exceeded the distance, the force of the ejection was
most remarkable. Are those who assert that Nature
knows no humour altogether justified ?

The examples so far cited and explained will
serve as a guide to those who desire to unravel the
secrets of the loves of the flowers. The bee we
have seen to play the part of fertiliser, so that upon
her action has depended the production of seed in
those plants which have lost the power of self-ferti¬
lisation ; but this is only one aspect of her work, for
she, the unconscious instrument, in a Hand unseen,
has been made to suffuse the landscape with colour,
and strew the path of man with the beauties of the
floral world. The homely garb of self-fertilised and
anemophilous flowers, such as those of the chick-
weed, the nettle, and the dock, indicates what all
would have been without insect action. In many
genera, the species present the greatest diversity with
regard to the size and beauty of their blossoms— e.g.,
Epilobium angustifolium has handsome and con¬
spicuous flowers, disposed in dense racemes, and
which, being proterandrous, are absolutely dependent.
In Epilobium parviflorum , or palustre, the flowers are
small and solitary, while they are capable of setting
seed by themselves, for their anthers and stigmas are
mature contemporaneously. So with the geranium
family, the different species indicating by the sizes of
their flowers how far they need insect help : the large
Geranium pratense, impotent without insects; the
small pusillum, generally self-fertilised. The reason

BEES AS FLORISTS.

321

is apparent: all variations which render the blossoms
more attractive, either by scent, colour, size of
corolla, or quantity of nectar, make the insect visit
more sure, and therefore the production of seed more
likely. Thus, the conspicuous blossoms secure de¬
scendants which inherit the special variations of their
parents, and so, generation after generation, we have
selection in favour of conspicuous flowers, where
insects are at work. Their appreciation of colour,
because it has brought the blossom possessing it more
immediately into their view, and more surely under
their attention, has enabled them, through the ages,
to be preparing the specimens upon which man now
operates; he taking up the work where they have
left it, selecting, inoculating, and hybridising, according
to his own rules of taste, and developing a beauty which
insects alone could never have evolved. His are the
finishing touches, his the apparent effects ; yet no less
is it true, that the results of his floriculture would
never have been attainable without insect helpers.
It is equally certain, that the beautiful perfume, and
the nectar also, are, in their present development,
the outcome of repeated insect selection ; and here, it
seems to me, we get an inkling of a deep mystery :
Why is life, in all its forms so dependent upon the
fusion of two individual elements ? Is it not, that
thus the doorway of progress has been opened ? If
each alone had reproduced, itself all-in-all, advance
would have been impossible ; the insect and human
florists and pomologists, like the improvers of animal
races, would have had no platform for their opera¬
tion, and, not only the forms of life, but life itself,

2 C

322

BEES AND BEE-KEEPING.

would have been stereotyped unalterably, ever me¬
chanically giving repetition to identical phenomena.

A new consideration now awaits us : Bees are not
only florists—they are fruit-producers; our orchard
and fruit crops, and leguminous seeds, constituting
together no inconsiderable fraction of human food,
are very largely dependent upon insect agency,
and the fee paid for professional attendance on the
part of the little inoculator is nectar. Let us take, as
an example, the apple, a fruit which, from a utilitarian

A, Blossom (Natural Size)— s, Stigmas; a, Anthers; p. Petal; ca. Calyx;
s', Sepal; d, Dissepiment. B, Section through partly developed Fruit—
f, f, Fertilised Carpels ; u, Unfertilised ditto.

point of view, has, in this country, no equal. Its
pretty blossom carries five stigmas, three of which
remain in the section A, Fig. 69; to each stigma
belongs a dissepiment, or division, of the compounded
ovary constituting the core of the fruit. The stigma
comes to maturity before the anthers, as in Scrophu-
lana nodosa. Bees seeking nectar get dusted com¬
pletely, and then transfer the granules to the stigmas
of neighbouring blossoms, while they are constantly
at work in packing the excess into their corbiculse.

BEES AS FRUIT-PRODUCERS.

3 2 3

And here let us remark in a parenthesis that the
multitude of pollen granules furnished by entomo-
philous plants, although usually less than in the case
of the anemophilous, is, nevertheless, enormous; the
single paeony, e.g., yielding about three and a half
millions per flower, while the number of granules
actually utilised is measured by the number of the.
ovules. In the curious cleistogamous flowers produced
occasionally or regularly by not a few plants, and which
do not open at all, or only in part—as the scentless
and small autumnal blossoms of the violet—there is
no repast for the insect, of nectar there is none,
and not a granule of pollen to spare ; for the anthers
and stigma, especially the former, are extremely small.
Yet self-fertilisation is completed, and seeds are
abundantly furnished, for all causes of waste are
avoided. But to return.

The apple, as its blossom indicates, is, strictly, a
fusion of five fruits into one—hence called pseudo-
syncarpous—and demands, for its production in per¬
fection, no less than five independent fertilisations.
If none are effected, the calyx, which really forms
the flesh of the fruit, instead of swelling, dries, and
soon drops. An apple often develops, however, though
imperfectly, if four only of the stigmas have been
pollen dusted, but it rarely hangs long enough to
ripen, the first severe storm sending it to the pigs
as a windfall. I had 200 apples, that had dropped
during a gale, gathered promiscuously for a lecture
illustration, and the cause of falling, in every case but
eight, was traceable to imperfect fertilisation. These
fruits may be generally known by a deformity ; one part

2 C 2

324

BEES AND BEE-KEEPING.

has failed to grow, because there has been no diver¬
sion of nutrition towards it. Cutting it across with
a knife, we find its hollow cheek lies opposite
the unfertilised dissepiment ( u , B), containing only
shrivelled pips. Pears are less impatient of imper¬
fect insect action than apples, though the structure
of the flower is the same. Amongst small fruits,
gooseberries are proterandrous, and absolutely de¬
pendent on insects. The failure of this crop is not
so uniformly the result, as some suppose, of frost

Fig. 70.— Raspberry (Rubus idaeus, Order Rosacece ) being Fertilised, and
Section op same.

A, Flower (Magnified Twice)— p, p, Petals; a, a, Anthers; s, Stigma; no,
Nectary Openings ; nc, Nectar Cells ; D, Drupels. B, Section through Core,
or Torus (C) and Drupels (V)—ud, Unfertilised Drupel; ws, Withered Stigma ;
wa, Withered Anther.

(this browning the exposed fruits) ; cold weather at
the critical time, keeping bees within,. often being
the chief cause, and showing itself in the dropping
of the open flowers from the protected branches. It
is here significant, that currants, which ripen their
pistils and anthers simultaneously, are said to be less
tender than gooseberries.

BEES AS FRUIT-PRODUCERS.

325

The raspberry gives the bee good return for its
work by yielding honey of excellent quality, and the
whole arrangement of its inflorescence points to an
effort to secure crossing. The petals ( p , p , A, Fig. 70)
are small, and widely placed, while within them are
disposed about ninety anthers ( a , a), on longer and
shorter filaments, which are set back, away from the
stigmas (j), one of which is carried by each of the
sixty or seventy drupels (D) making up the raspberry.
Examining a flower with a hand magnifier, we find
a circle of glistening dots (no) upon the receptacle,
and between the anthers and drupels. These dots
consist of nectar, furnished by secreting cells ( nc)
beneath. The bee alights upon the only solid resting-
place, the drupels, and applies her tongue rapidly to
dot after dot, revolving her body during the operation,
by which she gets dusted, on one side and beneath,
with pollen. Passing to the next blossom, she repeats
the operation, commonly with a difference which is
of primal importance : she revolves in the opposite
direction, by which she brings into play new muscles,
and rests those of the side which have just been exer¬
cised. The result is evident: the pollen acquired
on the previous visit is applied to the numerous stig-
matic faces waiting to receive it, which, as we have
so often seen, again secure crossing. Each seed thus
fertilised is soon surrounded by the luscious envelope
which protects the seed from injury, and makes the
manufacture of raspberry jam a possibility. These
rounded, red masses are never formed unless fertilisa¬
tion has taken place, neither ripening nor growth
being possible in its absence. When the season

326 BEES AND BEE-KEEPING.

is closing, the raspberry frequently fails in developing
some, or many, of its drupels; they remain green
and shrunken, for hive bees are loth to venture abroad,
and wild ones are dying off, or seeking, in the case
of the females, winter quarters. Some complain that
bees eat fruit, a charge which need not be rebutted ;
but it is for the bee-keeper to proclaim that, while
they gather nectar for themselves, and also for the
benefit of their master, they confer a greater boon
on the fruit-grower, for they really give him his crop
in return. The flowers of the blackberry (Rubus fruti-
cosus ) are similar in structure, and the explanation
given fully applies to them.

If we look at a strawberry, which is of a similar
type to the foregoing, we find a vast number of
(popularly) seeds (really achenia') studding its sur¬
face. Every one of these possessed a style and
stigma, as at s, s, A, Fig. 71, and has had pollen
conveyed to it by the action of insects, bees mainly.
When the bee settles, she, in her circular walk, rubs
from her body on to the stigmas, pollen brought
from another flower, as in the raspberry, for the
stigmas are receptive before the anthers have begun
to dehisce. The fertilisation, as before, determines
nutrition to the part, and the flower-stalk, which
forms the strawberry, becomes a luscious parenchyma.
But if any stigmas remain unpollinated, no develop¬
ment occurs at that spot, and here the strawberry
continues (as at u, B) hard, shrunken, and green, even
when the fertilised portion is fully ripe. We must
all again and again have seen illustrations of this,
from which we learn, that every strawberry requires

BEES AS FRUIT-PRODUCERS.

327

from 100 to 200, or even 300, distinct fertilisations
for its perfect production.

It would be unwise to omit a practical matter in
this connection. There is a tendency to a separation
of the sexes in the cultivated strawberry, which,
Darwin observes, “ is far more strongly marked in
the United States than in Europe; ” and growers
would do well to note, that plants bearing unusually
large blossoms are frequently tending to become male,
and produce few fruits; while those of the same

Fig. 71.— Strawberry (Fragaria vesca, Order Rosacece), Partly and
Fully Grown.

A, Strawberry, Earlier Stage—a, Anther ; s, s, Stigmas ; p, Petal. B, Section of
Mature Strawberry—a', Withered Anther : f, Fertilised Achenium (popularly
Seed); u, Unfertilised ditto; s', Withered Stigma.

variety, and under the same treatment, that produce
small blossoms, are tending to become female, and are
abundant bearers, while they yield few runners.
Without care in selecting, the numerous runners of
the former would ultimately supplant the female
forms, and so ruin the stock for economic purposes.
Lecturing, some while since, to several of the largest
growers of strawberries in the kingdom, I found all
quite unaware of this fact—at least, on its scientific

328

BEES AND BEE-KEEPING.

side—although in my own small beds the differences
had been sufficiently conspicuous.

We have yet to apply the numerous facts and
natural laws we have considered to Practical Apiculture,
which has utilities beyond those generally supposed;
and we can now see the wisdom of Girard’s remark,
that “ all money thrown out of the window, in en¬
couraging apiculture, will come in again by the door,
with heavy interest.” My sketch, which does not
cover the ground, but yet, I hope, dots it with illus¬
trations that may illumine the rest, must be regarded
as completed. And it leaves us here. The bee, with
all its wonderfulness, is only one wheel within many : she
takes to truly give, for seeds, flowers, and fruits, follow
in her train. Her honey is but a fraction of the
results of her labours. Man has had tiny helpers that
he knew not of. While he, for seasons, has selected
and hybridised, they, for ages, have, with their
little powers, toiled along, perpetuating every move¬
ment of the world of flowers towards the beautiful.
Flowers, yours is equal wonder and equal praise;
for dimly through you both I see that the praise is
not yours at all, saying with Tennyson :

“Flower in the crannied wall,

I pluck you out of the crannies;—

Hold you here, root and all, in my hand,

Little flqwer;—but if I could understand
What you are, root and all, and all in all,

I should know what God and man is.”

INDEX.

Abdomen, rings of, 31
Abdominal plates, 153
Abnormal bees, 208
Accouplement in confinement im¬
possible, 206
Air sacs, 35

sacs filled by flight, 148
Albino drones, 117
Anaphis, 44
Andrenidae, 8

tongue of, 100
Angles of comb, 164
Anguiculi, 124
Antennae, 20, 30, 103
cleaner, 129
of catasetum, 319
Anthers, 252

appendages of, 293
hiding, 285

movements of, 281, 283, 296
on elastic filaments, 290
retrorse, 314

Ants fond of aphide honey, 276
Aphide honey, 270
Aphides, 271

dissection of, 274

eggs of, 273

endure cold, 273

formed by interior budding,

273

how to reduce, in number, 276
number of progeny of, 274
number of species of, 272
ovaries of, 274

Aphidius rosae, 278
Apidae, 8

Appendicular gland, 224
gland intracellular, 225
Apple blossom, its structure, 322
needs five fertilisations, 323
Appliances too numerous, 5
Arbutus Unedo, 295
Armor copulatrix, 202
Artificial pollen, 126
Ascending and descending flight,
143

Aucuba japonica, 260
Aura seminalis, 236
Automatic movements of pulvillus,
127

Backward flight, 141
Barberry, 297
Barbs of sting, 186
Bee, antennae of, 20, 30, 103
Bees as fertilisers, 277
as florists, 320
as fruit-producers, 322
blood of, 81
brain of, 52, 55
carry pollen of one kind, 319
chrysalis of, 23, 238
claws of, 124
colour sense of, 116, 321
decorated by orchids, 317
digestive system of, 57
gather aphide honey after rain,
276

330

INDEX.

Bees, genera of, 13
glands of, 76
grubs of, 19, 23, 238
hearing of, 107
hermaphrodite, 208
how to free, from hairs, 121
humble, 12

not always able to fly, 146
not exceptionally framed, 246
number of species of, 8
nurses, peculiarities of, 19, 81
ocelli of, 116
rose cutters, 9
silk of, 21, 73, 175, 241
thorax of, 120, 238
unloading nectar, 18, 67
unloading pollen, 18, 130
voice of, 149

void faeces on the wing, j 48
wallring on glass, 125
Blackberry, 326
Bombi, 12

Bombus muscorum, 13
Bouton, or spoon of tongue, 95
Bowel blind in larva, 64
casting, 21, 241, 243
how developed in larva, 63
Brain, dissection of, 52

of insects compared, 54
size of, 55

Breathing apparatus, 33, 149
Broken combs, treatment of, 168
Brood, food of, 81
weaning of, 82
Buckwheat, 254, 300
Bursa copulatrix, 229
Buzzing, 149

Calyx, 251

Cane sugar converted into grape
sugar, 100

Cappings do not touch honey, 174
imperfect, 175
of drone cells, 180
Catasetum, 319
Cell, exuvium left in, 24

making, how started, i6r
of queen, 27, 172, 243
of queen contains cast skins,

Cells impervious to dyes, 173
normally circular, 170
number of, to lib. wax, 171
of comb unequal, 167
of wasps, 171
rhombic bases of, 161
sizes of, 176

Central and side ducts of ligula, 95
duct of ligula, size of, 96
Cephalic ganglia, 52
Chitine, 30
Chorion of egg, 230
Chrysalis condition, 23, 238
development of, 240
Chyle stomach, 60, 71
Circulation in bee, 37
Classification of Animal World, 7
Claws of bees, 124
Cleistogamous flowers, 249, 323
Clover, Dutch, does not seed under
a net, 308

Coccinella enemy of aphis, 276
Cocoon, 2r, 73, 175, 241
Colon, 62

Colour sense in bees, 116, 321
Comb, angles of, 164, 167

broken, treatment of, 168
building, how commenced, 161
coloured by breeding, 181
economises space, 169
Combs illustrated, 17

meet hygienic requirements,
182

model of, to build, 163
not mathematically correct, 166
partly constructed of old
material, 172
strength of, 179
structure of, 28, 163
Commissural nerve fibres, 48
Composite, 302
Compound eyes, 111
Conoid hairs of antennae, 106
Corbicula, 131
Corolla, 251
Coxa, or hip, 123
Crucifer®, 313
Cucurbitaceae, 259
Cynipidae (some) enemies of aphis,
277

INDEX.

Daisy, 305
Dandelion, 305
Darts of sting, 185
Delphinium, 282

Development of drone organs, 200
of larva, 19, 239
of sting, 196
of wings, 134

Diaphragm of abdomen, 39
Digestion similar in all cases, 58
Digestive system, 57
Dimorphic flowers, 297
Dioecious plants, 260
Discovery of the source of wax, 152
Dissection, methods of, 74
of eye, 112
Dorsal vessel, 37
Drone, antenna of, 109
bee, 27

cells, girdering of, 180
cells, sealing of, 176
decapitated, living, 49
egg, 238

flight, why sonorous, 149
organs of, 198

organs of, how to evert, 204
organs of, immense, 199
sealing, theory regarding, 181
spermatic vesicles of, 200
without wax glands, 154
Drones, abnormal, 207
albino, 117

destroyed in autumn, 211
eyeless, 117
have no father, 223
not used as heat-producers, 209
why fleet, 206
why many produced, 210
Ductus ejaculatorius, 202
Dufour theory, 68

Egg apparatus in plants, 256

artificial impregnation of, 233
chorion of, 230
laying, 25, 237
micropyle of, 231
spermatozoa within, 233
Eggs in ovarian tubes, 215

Eggs, number of, laid by each queen,
26, 228

total weight from, of one
"3

dissection, 120
Embryo sac, 254
Endo-skeleton, 88
Epilobium angustifolium, 287
Epipharynx, 90
Ericaceae, 289
Erica Tetralix, 292
Evolution of sexes, periods of, 244
Experiment illustrating flight, 140
of Huber on origin of wax, 158
on cell formation, 170
Extra-floral nectaries, 264
Extravasation of wings in chrysalis,

Exuvium left in cell, 24, 243
Eye cleaning brush, 128
compound, 111
method of examining, 112
retains traces of its cell origin,
119

Eyeless drone, 117

of the sexes numbered and
compared, 118
simple, 116

Embeddi

ling for

Faeces, why voided on the wing,
148

Feeding brood in cells, 101
groove, 86
Feelers of sting, 191
Feeling hairs, 31, 109
Fertile worker food, 82
workers, 223

Fertilisation impossible in confine¬
ment, 206
of flowers, 255

Swammerdam’s theory regard¬
ing, 236

Fertilising pouch, 229, 234
Festoons of wax workers, 160
Flapping moves the bee onward,
139

Flight, ascending and descending,

experiment upon, 140

332

INDEX.

Flight of drone, why sonorous, 149
rate of, 138

Flowers preventing self-fertilisa¬
tion, 248
sleeping, 305

why they supply insects, 247
Flying backwards, 138, 141
Food of larva, 19, 81
Foot, how detached, 127
pulvillus of, 125
Foraging bees, 19
Form of queen cell, 17, 173
French bean, 308

Ganglia, chain of, 45
number of, 50

supplying legs and wings, 51
Gastric glands, 61
teeth, 61

Genders, mixed, 208
Gland, appendicular, of queen, 216
gastric, 61

innominate, of spermathecal
duct, 227
intercellular, 79
intracellular, 77
lubricating, of sting, 192
mucous, of drone, 200
of colon, 62

olfactory, of Wolff, 43, 79
poison, of sting, 188
wax, 155

Glands, Nos. 1,2,3 and 4> uses of, 80
Nos. 2 and 4 digestive, 81
Nos. 2 and 3, position of ducts
of, 86, 88
Golden rod, 303
Gooseberries, 324

Graphic method of obtaining wing
rate, 145

Grubs, 19, 23, 238

Hairs encased in wax, 156
uses of, 31, no

Haviland, Mr., on drone produc¬
tion, 210

Hazel nut, inflorescence of, 261
Head, why wedge shaped, 10 r

Hearing of bees proved by observa¬
tion, 108
organs, 107
Heart ot bee, 37
Heather, 292
Hermaphrodite bees, 208
flowers, 281

Hexagonal facets of compound eye
caused by interference, 112
form of cells caused by inter¬
ference, 170

Hickson, Dr., on eyes of insects,

Hive bee, economy of, 15
Homologies between drone and
queen, 200

Honey and nectar, difference be¬
tween, 100, 263
bee unloading, 18, 67, 230
cappings imperfect, 174
dew, 270
poisonous, 289
sac, 18, 60

Hooklets of wing, 137
Huber, Francois, on wax, 152

error of, with regard to queen
cocoon, 241

experiment by, on source of
wax, 158

Humble bees, nest of, 12
Humming, 149
Hunter, Dr., on wax, 152
Huxley on multiplication of aphis,
275

Hyacinth, nectaries of, 269
Hypodermis, 30
Hypo-pharyngeal plate, 77

Imago, 24, 29
In-breeding prevented, 206
Inferiority of queen to worker, 55,

213

Insects and plants mutually de¬
pendent, 251

Intelligence measured by brain, 55
Interference makes cells hexagonal,
J70

Intestines, large and small, 62
Invagination explained, 34, 63
Italian bee, 222

INDEX.

333

J-

Jelly, royal, 82, 243

Kalmia latifolia, 289
Knotted figwort, 284

Labial palpi, 91
Labiatae, 309
Labium, 91
Labrum, 90

Ladybird enemy of aphis, 276
Lamellae of colon, 62
Lamium album. 309
Larva, bowel of, how developed, 63
casts bowel, 21, 241, 243
development, 19, 239
food of, 81
in egg, 64
moulting, 20, 243
sealing of, 21, 180
weight of, 20, 50
Legs, antenna cleaner of, 129
as tool bearers, 127
eyebrush of, 128
muscles of, 123
of queen and drone, 132
pollen basket of, 131
use of spur of, 130
velum of, 128
Leguminosae, 305

Leuwenhoek’s experiment on sight
in insects, 115

Ligula, artificial distension of, 99
central and side ducts of, 95
rod of, 97

rod of, thrown out by blood, 98
structure of, 92
Linum grandiflorum, 299
Lubricating gland of sting, 192
Lythrum salicaria, 300

Malpighian vessels, 61
Mandibles, 90, 169
Maraldi’s reputed measurement
mythic, 165

Marey on wing movements, 146
Mastication of wax scales, 157

Maxillae, 91, 94
Maxillaiy palpi, 92
Megachile centuncularis, 9
Membrane of ligula, 95
Mentum, 91

Meso-cephalic pillars, 89
Mesophragma, 88
Metatarsus of third leg, 131
Micropylar aperture of seed, 258
Micropyle of egg, 231
Microscope, dissecting, 74
Microscopic dissection of brain, 52
images produced by cornea of
bee. 115

Midrib of comb, 161
Monoecious plants, 260
Mosaic vision, 115
Moulting, 20

Mounting spermatozoa for micro¬
scope, 201
Muscles of legs, 123
of outer jaw, 43
of stomach-mouth, 67
protractor linguae, 9
Muscular contraction, 41
structure of sting, 187
system, 40

Nature, unity of, 12
Nectar, conditions favourable to its
secretion, 264

converted into honey, 100, 263
protected by hairs, 282, 294, 313
unloading of, 18, 67
Nectaries of aphis, 271

of flowers, 264, 268, 325
position of, 265
Nerve of taste, 78
system, 45

system of larva and imago dif¬
ferent, 50

Nervures of wings, 135
Notch on front leg, 129
Nurse bees, 19

bees have gland No. 1 large, 81
Nymph, or chrysalis, 23, 240

Obturator muscle of spiracle, 150
Ocelli of bee, 116

334

INDEX.

(Esophagus, 59
Olfactory gland of Wolff, 79
Ommatidia, 112
Orchis Mono, 314
Ovaries of queen, 214
Ovary of plant, 252
Oviducts of queen, 215
structure of, 234

Ovule and seed not identical, 253
P.

Pace of bees, 138
Pap for grubs, 19, 81
Paraglossae, 96
Parthenogenesis, 220
Pedunculated bodies of Dujardin, 54
Pelargonium examined, 251
nectary of, 267
Pericardial cells, 39
Periods of evolution, 244
Peritracheal circulation, 40
Pincers of third leg, 130
Pistil, 252
Pisum sativum, 306
Pneumophyses, 202
Poison bag, 189
channels, 191
dries like gum, 192
sac and gland of old queens
useless, 195
Pollen, 16, 252
basket, 131
brushes, 131

carried of one kind only, 319
grains, contents of, 257
grains, form of, 256
held by hairs of thorax, 120
tube, 248, 257
unloading, 18, 130
varies in dimorphic and tri-
morphic plants, 299, 301
Pollinia of orchid, 316
Polygonum fagopyrum, 300
Primine and secundine, 254
Primula vulgaris, 298
Proterandrous blossoms, 280
Proterogynous flowers, 284
Protractor linguae muscle, 91
Pulvillus, automatic movement of,
127

of foot, 125

Pumping action of sting, 189
Purple loosestrife, 300
Pylorus, 61

Queen, antenna of, 105
attendants, 26, 85
bee, 25, 212
bee, egg laying, 25, 234
bee not a ruler, 24
casts bowel against side of cell,
2 43

cell, 27, 243

cell constructed of old material,

cells, why thick, 173

cocoon of, 241

dejections of, 84

drone breeder, 227

egg production immense, 26

fecundity of, 83

food, of, 83

has no wax glands, 154
inferior to worker, 55, 213
marital flight of, 222
mating age, 221
organs of, 212
ovaries of, 214
spermatheca of, 216
sting of, 188
tongue of, 99
urinary tubes large, 84
weight of, 83
weight of eggs of, 83
Queens drone breeders, 222

drone breeders through paraly¬
sis, 236

have not gland No. i, 80
if frozen become drone breeders,
235

rapidly developed, 244
sting of, 194
variations in, 80
vary, 239

Raspberry, 324

Rate of wing vibration, 144

Reflex action, 48

Rod of ligula, 97

Rosebay willow herb, 287

INDEX.

335

Rose cutting bee, 9
Royal jelly, 82

Saccharine matter the source of
wax, 159
Sainfoin, 308

Saliva pumped out by bee in suck-

Salivary glands, discovery of, 72
glands, uses of, 80
valve, 79, 91, 101
Salvia glutinosa, 313
officinalis, 311
patens and fulgens, 312
Scarlet runner, 308
Schiemenz on stomach-mouth, 68
Scrophularia nodosa, 284
Sealing over brood pervious to air,

22> 175

Self-fertilised flowers small, 320
Shower of aphide nectar, 272
Side ducts extravasated, reason for,
97

Simpson’s honey plant, 286
Sir J. Lubbock on bees not hear¬
ing, 107

Sizes of antenna of the sexes, 110
of cells, 176
Skeleton, external, 30
Skin casting, 20
Sleeping flowers, 305
Smell, experiments upon, r09

hollows of antenna, 106,109, ill
Soap-bubble experiment, 170
Spermatheca, 216

of unmated queen, 218
valve of, 219
Spermathecal duct, 225
sphincter, 225
Spermatophore, 203
Spermatozoa, 200
diseased, 232
killed by cold, 235
movements of, 219
number of, 226
within egg, 233
Spinning gland, 73
Spiracles, 33, 149

can be closed voluntarily, 148
number of, 36

Spur on second leg, use of, 130
Stemmata, or simple eyes, 116
Sticky hairs of Erica Tetralix,
292

hairs of Salvia glutinosa, 313
Stigma, 252

of pelargonium, 283
Sting, antidote for, 188
darts of, 185

development of, 196, 240

how it pierces, 186

levers of, 186

lubricating gland of, 192

mechanical perfection of, 193

muscular structure of, 187

necessity for, 196

of queen, 193

palpi, 191

poison of, 188

pump of, 189

reflex action of, 187

sheath of, 184

structure, 183

valves of, 189

when used, tom from worker,
194

Stomach-mouth, 60, 65
movements of, 66
structure of, 67
true use of, 69

Stomato-gastric nerve system, 55
Strawberry, 326

culture : a caution, 326
needs hundreds of fertilisations,
326

tends to be dioecious, 327
tree, 295

Strength of comb, 179
Sub-oesophageal ganglion, 45
Sucking large quantities of nectar,
„ 94

Sugar does not fully replace honey,

Swarm, behaviour of, 160

cluster, how held together, 123
Swarming, 28
Sympathetic system, 55
Syrphidae and aphis, 277
Syrup, thick, slowly taken, 101
System No. 1, No. 2, and No. 3,
gland, illustrated, 75

336

INDEX.

Tactile hairs, 93, no
Taste, sense of, 103
Tendons, 42

Theory of backward flight, 143
Thistle, 303, 304
Thorax of bees, 120

development of, 240
Tongue, how folded out of view, 99
of Andrenidse, how folded, 100
structure of, settled by experi¬
ment, 97

Trachea;, minuteness of, 35
moulted, 34
of wax glands, 156
spiral thread of, 33
Transition cells, 177
Trifoliums, 308
Trimorphic flowers, 300
Tropaeolum majus, 268, 280

Vasa deferentia, 200
Vegetable marrow, 259
Velum of front leg, 128
Vesicula seminalis, 261
Vestibule of trachea, 150
Vetch, 308

Viola tricolor, nectar cells of, 266
Voice of bees, 149

Wasps encouraged by figwort, 286

Wax at first liquid, 156
costly to bee, 171
glands, 155
not a ^460

not ct&fived from pollen, 158
of stares and of comb unlike,

, 156 •

plate pincers, 130
pockets, 154
scales, 152

scales, mastication of, 157
secreted by a solitary bee, 160
secreting cells, 155
secretion exhausting, 159
secretion, favourable condi¬
tions for, 160
white, 181

Wild bees, nest of, 10
Wind a plant fertiliser, 250
Windfall apples, 323
Wing cells, 135

vibration, rate of, 144
Wings, development of, 134
hooklets of, 137
of drone, worker, and queen,
compared, 138
why in two pairs, 136
Worker egg a misnomer, 238
ovaries of, 238
superior to queen, 213
Workers, fertile, 223

Xenophon and wild honey, 291

Catalogue of Practical Handbooks
Ptiblished by L. Uftcott Gill\ iyo,
Strand, London , W.C.

ANGLER, BOOK OP THE ALL-ROUND. A Comprehensive
Treatise on Angling in both Fresh and Salt Water. In Four Divisions,
as named below. By John Bickerdyke. With over 220 Engravings,
In cloth, price 5s. 6d., by post 6s. (A few copies of a Large Paper
Edition, bound in Roxburghe, price 25s.)

Angling - for Coarse Pish. Bottom Fishing, according to the
Methods in use on the Thames, Trent, Norfolk Broads, and elsewhere.
Illustrated. In paper, price Is., by post Is. 2d.

Angling for Pike. The most Approved Methods of Fishing
for Pike or Jack. Profusely Illustrated. In paper, price Is., by post
Is. 2d.; cloth, 2s. {uncut), by post 2s. 3d.

Angling for Game Fish. The Various Methods of Fishing for
Salmon; Moorland, Chalk-stream, and Thames Trout; Grayling and
Char. Well Illustrated. In paper, price Is. 6d., by post Is. 9d.

Angling in Salt Water. Sea Fishing with Bod and Line, from
the Shore, Piers, Jetties, Bocks, and from Boats; together with Some
Account of Hand.-Lining. Over 50 Engravings. In paper, price Is., by
post Is. 2d.; cloth, 2s. (uncut), by post 2s. 3d.

AQUARIA, BOOK OF. A Practical Guide to the Construction,
Arrangement, and Management of Fresh-water and Marine Aquaria;
containing Full Information as to the Plants, Weeds, Fish, Molluscs,
Insects, &e., How and Where to Obtain Them, and How to Keep Them
in Health. Illustrated. By Bev. Gregory C. Bateman, A.K.C., and
Beginald A. B. Bennett, B.A. In cloth gilt,price 5s. 6d., by post 5s. lOd.

AQUARIA, FRESHWATER s Their Construction, Arrangement,
Stocking, and Management. Fully Illustrated. By Bev. G. C. Bate¬
man, A.K.C. In cloth gilt, price 3s. 6d., by post 3s. lOd.

AQUARIA, MARINE : Their Construction, Arrangement, and Manage¬
ment. Fully Illustrated. By B. A. B. Bennett, B.A. In cloth gilt,
price 2s. 6d., by post 2s. 9d.

AUSTRALIA, SHALL I TRY? A Guide to the Australian
Colonies for the Emigrant Settler and Business Man. With two
Illustrations. By George Lacon James. In cloth gilt, price 3s. 6d.,
by post 3s. lOd.

AUTOGRAPH COLLECTING : A Practical Manual for Amateurs
and Historical Students, containing ample information on the Selec¬
tion and Arrangement of Autographs, the Detection of Forged
Specimens, &c., &c., to which are added numerous Facsimiles for
Study and Beference, and an extensive Valuation Table of Auto¬
graphs worth Collecting. By Henry T— Scott, M.D., L.B.C.P., &c.,
Bector of Swettenham, Cheshire. In leatherette gilt, price 7s. 6d., by
post 7s, lOd.

BAZAARS AND FANCY FAIRS : Their Organization and Manage-
ment. A Secretary’s Fade Mecum. By John Muir. In paper,
price Is., by post Is. 2d.

170 C 1/96

Published by L. Upcott Gill,

BEES AND BEE-KEEPING : Scientific and Practical. By F. R.
Cheshire, F.L.S., F.R.M.S., Lecturer on Apiculture at South Kensington.
In two vols., cloth gilt, price 16s., by post 16s. 8dt.

Vol. I., Scientific. A complete Treatise on the Anatomy and
Physiology of the Hive Bee. In cloth gilt, price 7s. 6 d., by post 7s. 10d.

Vol. II., Practical management of Bees. An Exhaustive
Treatise on Advanced Bee Culture. In cloth gilt, price 8s. 6 d., by
post 9s.

BEE-KEEPING, BOOK OP. A very practical and Complete Manual
on the Proper Management of Bees, especially written for Beginners
and Amateurs who have but a few Hives. Fully Illustrated. By
W. B. Webster, First-class Expert, B.B.K.A. In paper, price Is.,
by post Is. 2 d .; cloth, Is. 6 d., by post Is. 8 d.

BEGONIA CULTURE, for Amateurs and Professionals. Containing
Full Directions for the Successful Cultivation of the Begonia, under
Glass and in the Open Air. Illustrated. By B. C. Ravenscroft. In
paper, price Is., by post Is. 2 d.

BENT IRON WORK : A Practical Manual of Instruction for Amateurs
in the Art and Craft of Making and Ornamenting Light Articles in
imitation of the beautiful Mediaeval and Italian Wrought Iron Work.
By F. J. Erskine. Illustrated. In paper, price Is., by post Is. 2d.

BOAT BUILDING AND SAILING, PRACTICAL. Containing
Full Instructions for Designing and Building Punts, Skiffs, Canoes,
Sailing Boats, &c. Particulars of the most suitable Sailing Boats and
Yachts for Amateurs, and Instructions for their Proper Handling. Fully
Illustrated with Designs and Working Diagrams. By Adrian Neison,
C.E., Dixon Kemp, A.I.N.A., and G. Christopher Davies. In one vol.,
cloth gilt, price 7s. 6 d., by post 7s. lOd.

BOAT BUILDING FOR AMATEURS, PRACTICAL. Contain¬
ing Full Instructions for Designing and Building Punts, Skiffs, Canoes,
Sailing Boats, &c. Fully Hlustrated with Working Diagrams. By
Adrian Neison, C.E. Second Edition, Revised and Enlarged by Dixon
Kemp, Author of “Yacht Designing,” “A Manual of Yacht and Boat
Sailing,” &c. In cloth gilt, price 2s. 6 d., by post 2s. 9d.

BOAT SAILING FOR AMATEURS. Containing Particulars of
the most Suitable Sailing Boats and Yachts for Amateurs, and Instruc¬
tions for their Proper Handling, &e. Illustrated with numerous
Diagrams. By G. Christopher Davies. Second Edition, Revised and
Enlarged, and with several New Plans of Yachts. In cloth gilt, price 5s.,
by post 5s. 4 d.

BOOKBINDING FOR AMATEURS ; Being Descriptions of the
various Tools and Appliances Required, and Minute Instructions for
their Effective Use. By W. J. E. Crane. Illustrated with 156 Engrav¬
ings. In cloth gilt, price 2s. 6 d., by post 2s. 9 d.

BUNKUM ENTERTAINMENTS : A Collection of Original Laugh¬
able Skits on Conjuring, Physiognomy, Juggling, Performing Fleas,
Waxworks, Panorama, Phrenology, Phonograph, Second Sight,
Lightning Calculators, Ventriloquism, Spiritualism, &c., to which are
added Humorous Sketches, Whimsical Recitals, and Drawing-room
Comedies. In cloth, price 2s. 6 d., by post 2s. 9 d.

BUTTERFLIES, THE BOOK OF BRITISH: A Practical
Manual for Collectors and Naturalists. Splendidly Hlustrated through¬
out with very accurate Engravings of the Caterpillars, Chrysalids, and
Butterflies, both upper and under sides, from drawings by the Author
or direct from Nature. By W. J. Lucas, B.A. Price 3s. 6 d., by post 3s. 9d.

170 , Strand, London, W.C.

BUTTERFLY AND MOTH COLLECTING : Where to Search,
and What to Do. By G. E. Simms. Illustrated. In paper, price Is.,
by post Is. 2 d.

CACTUS CULTURE FOR AMATEURS: Being Descriptions of
the various Cactuses grown in this country; with Full and Practical
Instructions for their Successful Cultivation. By W. Watson, Assistant
Curator of the Royal Botanic Gardens, Kew. Profusely Illustrated.
In cloth gilt, price 5s., by post 5s. id.

CAGE BIRDS, DISEASES OF : Their Causes, Symptoms, and Treat,
ment. A Handbook for everyone who keeps a Bird. By Db. W. T.
Gbeene, F.Z.S. In paper, price Is., by post Is. 2d.

CANARY BOOK. The Breeding, Rearing, and Management of
all Varieties of Canaries and Canary Mules, and all other matters
connected with this Fancy. By Robebt L. Wallace. Third
Edition. In cloth gilt, price 5s., by post 5s. 4d.; with COLOURED
PLATES, 6s. 6 d., by post 6s. lOd.; and as follows:

General Management of Canaries. Cages and Cage-making
Breeding, Managing, Mule Breeding, Diseases and their Treatment,
Moulting, Pests, &c. Illustrated. In cloth, price 2s. 6 d., by post 2s. 9d.

Exhibition Canaries. Full Particulars of all the different
Varieties, their Points of Excellence, Preparing Birds for Exhibi¬
tion, Formation and Management of Canary Societies and Exhibitions.
Illustrated. In cloth, price 2s. 6d., by post 2s. 9 d.

CANARY, THE FET. With some Instructions as to its Purchase,
Diet, Toilette, Cage, and Sanitary Keeping. Written for the Guidance
of the Amateur. By W. H. Betts, Honorary Treasurer Cage Bird Club,
with Specially designed Illustrations by A. F. Lydon, G. R. Kennebley,
and Mobant Cox. 250 Copies (Numbered), printed on thick antique
paper, and bound in cloth. Crown Svo, price 5s., by post 5s. 3d.

CARD TRICKS, BOOK OF, for Drawing-room and Stage Entertain¬
ments by Amateurs ; with an exposure of Tricks as practised by Card
Sharpers and Swindlers. Numerous Illustrations. By Pboe. R. Ktjnaed.
In illustrated wrapper, price 2s. 6<£., by post 2s. 9d.

CATS, DOMESTIC OR FANCY: A Practical Treatise on their
Antiquity, Domestication, Varieties, Breeding, Management, Diseases
and Remedies, Exhibition and Judging. By John Jennings. Illus¬
trated. In cloth, price 2s. 6 d., by post 2s. 9 d.

CHRYSANTHEMUM CULTURE, for Amateurs and Professionals.
Containing Full Directions for the Successful Cultivation of the
Chrysanthemum for Exhibition and the Market. Illustrated. By
B. C. Ravenscboet. In paper, price Is., by post Is. 2d.

COINS, A GUIDE TO ENGLISH PATTERN, in Gold, Silver,
Copper, and Pewter, from Edward I. to Victoria, with their Value. By
the Rev. G. F. Cbowtheb, M.A. Illustrated. In silver cloth, with gilt
facsimiles of Coins, price 5s., by post 5s. 3d.

COINS OF GREAT BRITAIN AND IRELAND, A GUIDE
TO THE, in Gold, Silver and Copper, from the Earliest Period to the
Present Time, with their Value. By the late Colonel W. Stewabt
Thobbtjbn. With 32 Plates in Gold, Silver, Copper, &c. In cloth
gilt, price 7s. 6 d., by post 7s. lOd.

COLLIE, THE. Its History, Points, and Breeding. By Hugh Dalziel.
Illustrated with Coloured Frontispiece and Plates. In paper, price Is.,
by post Is. 2d .; cloth, 2s., by post 2s. 3d.

COLLIE STUD BOOK. Edited by Hugh Dalziel. Price 3s. 6 d. each,
by post 3s. 9d. each.

4 Published by L. Upcott Gill,

Vol. I., containing Pedigrees of 1308 of the best-known Dogs, traced
to their most remote known ancestors ; Show Record to Feb., 1890, &c.

Vol. II. Pedigrees of 795 Dogs, Show Record, &c.

Vol. III. Pedigrees of 786 Dogs, Show Record, &c.

COLUMBARIUM, MOORE'S. Reprinted Verbatim from the original
Edition of 1735, with a Brief Notice of the Author. By W. B. Tkgkt-
meier, F.Z.S., Member of the British Ornithologists’ Union. Price Is.,
by post Is. 2 d.

CONJURING, BOOK OP MODERN. A Practical Guide to Drawing,
room and Stage Magic for Amateurs. By Professor R. Kunard.
Illustrated. In illustrated wrapper, price 2s. 6d., by post 2s. 9 d.

COOKERY FOR AMATEURS; or, French Dishes for English Homes
of all Classes. Includes Simple Cookery, Middle-class Cookery, Superior
Cookery, Cookery for Invalids, and Breakfast and Luncheon Cookery.
By Madame Valerie. Second Edition. In paper, price Is., by post Is. 2d.

CRICKET. The History of a Hundred Centuries, as Written by Dr.
W. G. Grace, and Edited by W. Yardley. In paper , price 6 d., by
post Id.

CUCUMBER CULTURE FOR AMATEURS. Including also
Melons, Vegetable Marrows, and Gourds. Illustrated. By W. J. May.
In paper, price Is., by post Is. 2d.

CYCLIST’S ROUTE MAP of England and Wales. Shows clearly
all the Main, and most of the Cross, Roads, and the Distances between
the Chief Towns, as well as the Mileage from London. In addition
to this, Routes of Thirty of the most Interesting Tours are printed in
red. The map is mounted on linen, and is the fullest, handiest, and best
tourist’s map in the market. In cloth, price Is., by post Is. 2d.

DESIGNING, HARMONIC. Explaining a System whereby an
Endless Variety of Most Beautiful Designs suited to numberless
Manufactures may be obtained by Unskilled Persons from any Printed
Music. Illustrated by Numerous Explanatory Diagrams and Illus¬
trative Examples. By C. H. Wilkinson, author of “ Harmonious
Colouring.” With Introductory Chapter by John D. Macdonald,
M.D., F.R.S. Demy 4 to, price £2 2s. ' m tln the Press.

DOGS, BREAKING AND TRAINING : Being Concise Directions
for the proper education of Dogs, both for the Field and for Companions.
Second Edition. By “Pathfinder.” With Chapters by Hugh
Dalziel. Illustrated. In cloth gilt, price 6s. 6d., by post 6s. 10d.

DOGS, BRITISH, ANCIENT AND MODERN : Their Varieties,
History, and Characteristics. By Hugh Dalziel, assisted by Eminent
Fanciers. SECOND EDITION, Revised and Enlarged. Illustrated with
First-class COLOURED PLATES and full-page Engravings of Dogs of
the Day. This is the fullest work on the various breeds of dogs kept in
England. In three volumes, demy 8 vo, cloth gilt, price 10s. 6cL each, by
post 11s. Id. each.

Dogs Used in Field Sports. Containing Particulars of the
following among other Breeds : Greyhound, Irish Wolfhound, Blood-
hound, Foxhound, Harrier, Basset, Dachshund, Pointer, Setters,
Spaniels, and Retrievers. Seven Coloured Plates and 21 full-page
Engravings.

Dogs Useful to Man in other Work than Field Sports ;
House and Toy Dogs. Containing Particulars of the following,
among other Breeds: Collie, Bulldog, Mastiff, St. Bernards, Newfoundland,
Great Dane, Fox and all other Terriers, King Charles and Blenheim

170 , Strand, London, W.C.

Spaniels, Png, Pomeranian, Poodle, Italian Greyhound, Toy Dogs, &o.,
&e. Coloured Plates and full-page Engravings.

Practical Kennel Management : A Complete Treatise on all
Matters relating to the Proper Management of Dogs, whether kept for
the Show Bench, for the Field, or for Companions. Illustrated with
Coloured and numerous other Plates. [In the Press.

BOGS, DISEASES OP : Their Causes, Symptoms, and Treatment;
Modes of Administering Medicines ; Treatment in cases of Poisoning, &c.
For the use of Amateurs. By Hugh Dalziel. Third Edition. In
paper, price Is., by post Is. 2d. ; in cloth gilt, 2s., by post 2s. 3d.

ENTERTAINMENTS, AMATEUR, POR CHARITABLE AND
OTHER OBJECTS : How to Organize and Work them with Profit
and Success. By Robert Ganthony. In colov/red cover, price Is., by
post Is. 2d.

PANCY WORK SERIES, ARTISTIC. A Series of Illustrated
Manuals on Artistic and Popular Fancy Work of various kinds. Each
number is complete in itself, and issued at the uniform price of 6d., by
post 7d. Now ready— (1) Macrame Lace (Second Edition); (2) Patch-
work ; (3) Tatting; (4) Crewel Work; (5) Applique; (6) Fancy
Netting.

FERNS, THE BOOK OF CHOICE : for the Garden, Conservatory,
and Stove. Describing the best and most striking Ferns and Sela-
ginellas, and giving explicit directions for their Cultivation, the for-
mation of Rockeries, the arrangement of Ferneries, &c. By George
Schneider. With numerous Coloured Plates and other Illustrations.
In 3 vols., large post too. Cloth gilt, price £3 3s., by post <£3 6s.

FERNS, CHOICE BRITISH. Descriptive of the most beautiful
Variations from the common forms, and their Culture. By C. T. Druery,
F.L.S. Very accurate PLATES, and other Illustrations. In cloth gilt,
price 2s. 6d., by post 2s. 9d.

FERRETS AND FERRETING. Containing Instructions for the
Breeding, Management, and Working of Ferrets. Second Edition, Re¬
written and greatly Enlarged. Illustrated. In paper, price 6d., by
post 7d.

FERTILITY OF EGGS CERTIFICATE. These are Forms of
Guarantee given by the Sellers to the Buyers of Eggs for Hatching,
undertaking to refund value of any unfertile eggs, or to replace them
with good ones. Very valuable to sellers of eggs, as they induce
purchases. In boohs, with counterfoils, price 6d., by post 7d.

FIREWORK-MAKING POR AMATEURS. A complete, accurate,
and easily-understood work on Making Simple and High-class Fireworks.
By Dr. W. H. Browne, M.A. In paper, price 2s. 6d., by post 2s. 9d.

FISHERMAN, THE PRACTICAL. Dealing with the Natural
History, the Legendary Lore, the Capture of British Fresh-Water Fish,
and Tackle and Tackle-making. By J. H. Keene. In cloth, price
7s. 6d., by post, 8s.

FOREIGN BIRDS, FAVOURITE, for Cages and Aviaries. How to
Keep them in Health. Fully Illustrated. By W. T. Greene, M.A.,
M.D., F.Z.S., &e. In cloth, price 2s. 6d., by post 2s. 3d.

FOX TERRIER, THE. Its History, Points, Breeding, Rearing, Pre¬
paring, for Exhibition, and Coursing. By Hugh Dalziel. Illustrated
with Coloured Frontispiece and Plates. In paper, price Is., by post
Is. 2d. ; cloth, 2s., by post 2s. 3d.

Published by L. Ufcott Gill,

FOX TSEEIER STUB BOOK. Edited by Hugh Dalziel. Price
3s. 6 d. each., by post 3s. 9d. each.

Vol. I., containing Pedigrees of over 1400 of the best-known Dogs,
traced to their most remote known ancestors.

Vol. II. Pedigrees of 1544 Dogs, Show Record, &c.

Vol. III. Pedigrees of 1214 Dogs, Show Record, &c.

Vol. IV. Pedigrees of 1168 Dogs, Show Record, &e.

Vol. V. Pedigrees of 1662 Dogs, Show Record, &c.

FRETWORK AND MARQUETRY. A Practical Manual of
Instructions in the Art of Fret-cutting and Marquetry Work. Profusely
Illustrated. By D. Denning. In cloth, price 2s. 3d., by post 2s. lOd.

FRIESLAND MERES, A CRUISE ON THE. By Ernest R.
Suppling. Illustrated. In paper, price Is., by post Is. 2 d.

GAME AND GAME SHOOTING, NOTES ON. Grouse, Par¬
tridges, Pheasants, Hares, Rabbits, Quails, Woodcocks, Snipe, and
Rooks. By J. J. Manley. Illustrated. In cloth gilt, price 7s. 6d.,
by post 7s. IOcZ.

GARDENING, DICTIONARY OF. A Practical Encyclopedia of
Horticulture, for Amateurs and Professionals. Illustrated with 2440
Engravings. Edited by G. Nicholson, Curator of the Royal Botanic
Gardens, Kew; assisted by Prof. Trail, M.D., Rev. P. W. Myles,
B.A., F.L.S., W. Watson, J. Garrett, and other Specialists. In 4 uols.,
large post 4to. In cloth gilt, price £ 3, by post £3 3s.

GARDENING IN EGYPT. A Handbook of Gardening for Lower
Egypt. With a Calendar of Work for the different Months of the
Year. By Walter Draper, Fellow of the Royal Horticultural
Society; Director of Government Gardens at the Barrage, near Cairo.
In cloth, price 3s. 3d., by post 3s. 9 d.

GOAT, BOOK OF THE. Containing Full Particulars of the various
Breeds of Goats, and their Profitable Management. With many Plates.
By H. Stephen Holmes Pegler. Third Edition, with Engravings and
Coloured Frontispiece. In cloth gilt, price 4s. 3d., by post 4s. 10d.

GOAT-KEEPING FOR AMATEURS : Being the Practical Manage-
ment of Goats for Milking Purposes. Abridged from “ The Book of the
Goat.” Hlustrated. In paper, price Is., by post Is. 2 d.

GRAPE GROWING FOR AMATEURS. A Thoroughly Practical
Book on Successful Vine Culture. By E. Molyneux. Illustrated. In
paper, price Is., by post Is. 2 d.

GREENHOUSE MANAGEMENT FOR AMATEURS. The

Best Greenhouses and Frames, and How to Build and Heat them, Illus¬
trated Descriptions of the most suitable Plants, with general and
Special Cultural Directions, and all necessary information for the
Guidance of the Amateur. Second Edition, Revised and Enlarged.
Magnificently Illustrated. By W. J. May. In cloth gilt, price 5s.,
by post 5s. 4 d.

GREYHOUND, THE : Its History, Points, Breeding, Rearing, Training,
and Running. By Hugh Dalziel. With Coloured Frontispiece. In
cloth gilt, demy 8 vo, price 2s. 3d., by post 2s. 9d.

GUINEA PIG, THE, for Food, Fur, and Fancy. Illustrated with
Coloured Frontispiece and Engravings. An exhaustive book on the
Varieties of the Guinea Pig, and its Management. By C. Cumberland,
F.Z.S. In cloth gilt, price 2s. 3d., by post 2s. 9d.

170 , Strand, London, W.C.

HAND CAMERA MANUAL, THE. A Practical Handbook on all
Matters connected with the Use of the Hand Camera in Photography.
Illustrated. By W. D. Weleord. Third Edition. Price Is., by post Is. 2d.

HANDWRITING, CHARACTER INDICATED BY. With Illus¬
trations in Support of the Theories advanced taken from Autograph
Letters of Statesmen, Lawyers, Soldiers, Ecclesiastics, Authors, Poets,
Musicians, Actors, and other persons. Second Edition. By B. Baughan.
In cloth gilt , price 2s. 6 d., by post 2s. 9d.

HARDY PERENNIALS and Old-fashioned Garden Flowers. Descrip¬
tions, alphabetically arranged, of the most desirable Plants for Borders,
Bockeries, and Shrubberies, including Foliage as well as Flowering
Plants. Profusely Illustrated. By J. Wood. In cloth, price 5s., by
post 5s. 4 d.

HAWK MOTHS, BOOK OP BRITISH. A Popular and Practical
Manual for all Lepidopterists. Copiously illustrated in black and white
from the Author’s own exquisite Drawings from Nature. By W. J.
Lucas, B.A. In cloth, price 3s. 6d., by post 3s. 9 d.

HOME MEDICINE AND SURGERY: A Dictionary of Diseases
and Accidents, and their proper Home Treatment, For Family Use. By
W. J. Mackenzie, M.D. Illustrated. Incloth, price 2s. 6d., by post 2s. 9d.

HORSE-KEEPER, THE PRACTICAL. By George Fleming, C.B.,
LL.D., F.B.C.V.S., late Principal Veterinary Surgeon to the British
Army, and Ex-President of the Boyal College of Veterinary Surgeons.
In cloth, price 3s. 6d., by post 3s. lOd.

HORSE-KEEPING TOR AMATEURS. A Practical Manual on
the Management of Horses, for the guidance of those who keep one or
two for their personal use. By Fox Bussell. In paper, price Is.,
by post Is. 2 d.; cloth, 2s., by post 2s. 3d.

HORSES, DISEASES OP: Their Causes, Symptoms, and Treatment.
For the use of Amateurs. By Hugh Dalziel. In paper, price Is.,
by post Is. 2d.; cloth 2s., by post 2s. 3d.

INLAND WATERING PLACES. A Description of the Spas of
Great Britain and Ireland, their Mineral Waters, and their Medicinal
Value, and the attractions which they offer to Invalids and other Visitors.
Profusely illustrated. A Companion V olume to ‘ 1 Seaside Watering Places.”
In cloth, price 2s. 6d., by post 2s. lOd.

JOURNALISM, PRACTICAL : How to Enter Thereon and Succeed.
A book for all who think of “ writing for the Press.” By John Dawson.
In cloth gilt, price 2s. 6d., by post 2s. 9d.

LAYING HENS, HOW TO KEEP and to Bear Chickens in Large
or Small Numbers, in Absolute Confinement, with Perfect Success. By
Major G. F. Morant. In paper, price 6d., by post 7d.

LIBRARY MANUAL, THE. A Guide to the Formation of a Library,
and the Values of Bare and Standard Books. By J. H. Slater,
Barrister-at-Law. Third Edition. Bevised and Greatly Enlarged. In
cloth gilt, price 7s. 6d., by post 7s. lOd.

MAGIC LANTERNS, MODERN, A Guide to the Management
of the Optical Lantern, for the Use of Entertainers, Lecturers,
Photographers, Teachers, and others. By B. Child Bayley. In
paper, price Is., by post Is. 2d.

MICE, FANCY : Their Varieties, Management, and Breeding. Ee-issue,
with Criticisms and Notes by Dr. Carter Blake. Illustrated. In
paper, price 6d., by post 7d.

Published by L. Upcott Gill,

MILLINERY, HANDBOOK OF. A Practical Manual of Instruction
for Ladies. Illustrated. By Mme. Rose®, Court Milliner, Principal of
the School of Millinery. In paper, price Is., by post Is. 2 d.

MODEL YACHTS AND BOATS : Their Designing, Making, and
Sailing. Illustrated with 118 Designs and Working Diagrams. A
splendid book for boys and others interested in making and rigging toy
boats for sailing. It is the best book on the subject now published. By
J. du Y. Grosvenor. In leatherette, price 5s., by post 5s. 3d.

MONKEYS, PET, and How to Manage Them. Illustrated. By Aethub
Patteeson. In cloth gilt, price 2s. 6d., by post 2s. 9d.

MUSHROOM CULTURE FOR AMATEURS. With Full Directions
for Successful Growth in Houses, Sheds, Cellars, and Pots, on Shelves,
and Out of Doors. Illustrated. By W. J. May. In paper, price Is, by
post Is. 2d.

NATURAL HISTORY SKETCHES among the Carnivora—Wild and
Domesticated; with Observations on their Habits and Mental Faculties.
By Arthub Nicols, F.G.S., F.R.G.S. Illustrated. In cloth gilt,
price 2s. 6d., by post 2s. 9d.

NEEDLEWORK, DICTIONARY OF. An Encyclopaedia of Artistic,
Plain, and Fancy Needlework ; Plain, practical, complete, and
magnificently Illustrated. By S. F. A. Caulfeild and B. C. Saward.
Accepted by H.M. the Queen, H.R.H. the Princess of Wales, H.R.H. the
Duchess of Edinburgh, H.R.H. the Duchess of Connaught, and H.R.H.
the Duchess of Albany. Dedicated by special permission to H.R.H.
Princess Louise, Marchioness of Lome. In demy 4fo, 528pp., 829
Illustrations, extra cloth gilt, plain edges, cushioned bevelled boards, price
21s., by post 22s.; with COLOURED PLATES, elegant satin brocade
cloth binding, and coloured edges, 31s. 6d., by post 32s. 6d.

ORCHIDS : Their Culture and Management, with Descriptions
of all the Kinds in General Cultivation. Illustrated by Coloured
Plates and Engravings. By W. Watson, Assistant-Curator,
Royal Botanic Gardens, Kew; Assisted by W. Bean, Foreman, Royal
Gardens, Kew. Second Edition, Revised and with Extra Plates. In
cloth gilt and gilt edges, price £1 Is., by post £1 Is. lOd.

PAINTERS AND THEIR WORKS. A Work of the Greatest
■ Value to Collectors and such as are interested in Art, as it gives,
besides Biographical Sketches of all the Artists of Repute (not now
living) from the 13th Century to the present date, the Market Value
of the Principal Works Painted by Them, with Full Descriptions of
Same. In 2 vols., cloth, price 31s. 6<Z., by post 32s.

PAINTING, DECORATIVE. A practical Handbook on Painting and
Etching upon Textiles, Pottery, Porcelain, Paper, Vellum, Leather, Glass,
Wood, Stone, Metals, and Plaster, for the Decoration of our Homes. By
B. C. Saward. In cloth, price 5s., by post 5s. 4d.

PARCEL POST DISPATCH BOOK (registered). An invaluable book
for all who send parcels by post. Provides Address Labels, Certificate
of Posting, and Record of Parcels Dispatched. By the use of this book
parcels are insured against loss or damage to the extent of £2.
Authorized by the Post Office. Price Is., by post Is. 2 d., for 100
parcels; larger sizes if required.

PARROT, THE GREY, and How to Treat it. By W. T. Greene,
M.D., M.A., F.Z.S., &e. Price Is., by post Is. 2d.

PARROTS, THE SPEAKING. The Art of Keeping and Breeding
the principal Talking Parrots in Confinement. By Dr. Karl Russ.

170 , Strand, London, W.C.

Illustrated with COLOUBED PLATES and Engravings. In cloth gilt,
price 5s., by post 5s. 4 d.

PATIENCE, GAMES OP, for one or more Players. How to Play
106 different Games of Patience. By Miss Whitmore Jones. Illus¬
trated. Series I., 39 games; Series II., 31 games; Series III., 33
games. Each Is., by post Is. 2d. The three bound together m cloth,
price 3s. 6d., by post 3s. lOd.

FEN PICTURES, and How to Draw Them. A Practical Handbook on
the various Methods of Illustrating in Black and White for “Process”
Engraving, with numerous Designs, Diagrams, and Sketches. By Eric
Meade. In cloth gilt, price 2s. 6d., by post 2s. 9d.

PERSPECTIVE, THE ESSENTIALS OP. With numerous
Illustrations drawn by the Author. By L. W. Miller, Principal of the
School of Industrial Art of the Pennsylvania Museum, Philadelphia.
This book is such a manual as has long been desired for the guidance
of art students and for self-instruction. The instructions are clearly
set forth, and the principles are vividly enforced by a large number
of attractive drawings. Price 6s. 6 d., by post 6s. lOd.

PHEASANT-KEEPING FOR AMATEURS. A Practical Hand-
book on the Breeding, Bearing, and General Management of Fancy
Pheasants in Confinement. By Geo. Horne. Fully Illustrated. In
cloth gilt, price 3s. 6c?., by post 3s. 9cZ.

PHOTOGRAPHY (MODERN) FOR AMATEURS. New and
Bevised Edition. By J. Eaton Fearn. In paper, price Is., by post
Is. 2d.

PIANOFORTES, TUNING AND REPAIRING. The Amateur’s
Guide to the Practical Management of a Piano without the intervention
of a Professional. By Charles Babbington. In paper, price 6 <?., by
post 6 id.

PICTURE-FRAME MAKING FOR AMATEURS. Being Practi-
cal Instructions in the Making of various kinds of Frames for Paintings,
Drawings, Photographs, and Engravings. Illustrated. By the Bev. J.
Lukin. Cheap Edition, in paper, price Is., by post Is. 2d.

PIG, BOOK OF THE. The Selection, Breeding, Feeding, and
Management of the Pig; the Treatment of its Diseases; the Curing
and Preserving of Hams, Bacon, and other Pork Foods; and other
information appertaining to Pork Farming. By Professor James Long.
Fully Illustrated with Portraits of Prize Pigs, Plans of Model Piggeries,
&c. In cloth gilt, price 10s. 6<?., by post 11s. Id.

PIG-KEEPING, PRACTICAL : A Manual for Amateurs, based on
Personal Experience in Breeding, Feeding, and Fattening; also in
Buying and Selling Pigs at Market Prices. By B. D. Garratt. In
paper, price Is., by post Is. 2d.

PIGEONS, FANCY. Containing Full Directions for the Breeding and
Management of Fancy Pigeons, and Descriptions of every known
Variety, together with all other information of interest or use to Pigeon
Fanciers. Third Edition, bringing the subject down to the present
time. 18 COLOUBED PLATES, and 22 other full-page Illustrations.
By J. C. Lyell. In cloth gilt, price. 10s. 6d., by post 10s. lOd.

PIGEON-KEEPING FOR AMATEURS. A complete Guide to the
Amateur Breeder of Domestic and Fancy Pigeons. By J. C. Lyell.
Illustrated. In cloth, price 2s. 6<?., by post 2s. 9d.

POKER BOOK, THE. How to Play Poker with Success. By B.
Guerndale. In paper, price Is., by post Is. 2d.

Published by L. Upcott Gill,

POLISHES AND STAINS POR WOODS : A Complete Guide to
Polishing Woodwork, with Directions for Staining, and Pull Information
for making the Stains, Polishes, &c., in the simplest and most satis¬
factory manner. By David Dunning. In paper, price Is., by post
Is. 2 d.

POOL, GAMES OP. Describing Various English and American
Pool Games, and giving the Buies in full. Illustrated. In paper,
price Is., by post Is. 2d.

POTTERY AND PORCELAIN, ENGLISH. A Guide for
Collectors. Handsomely Illustrated with Engravings of Specimen Pieces
and the Marks used by the different Makers. New Edition, Bevised
and Enlarged. By the Eev. E. A. Downman. In cloth gilt, price 5s.,
by post 5s. 4d.

POULTRY-KEEPING, POPULAR. A Practical and Complete
Guide to Breeding and Keeping Poultry for Eggs or for the Table. By
E. A. Mackenzie. Illustrated. In paper, price Is., by post Is. 2d.

POULTRY AND PIGEON DISEASES : Their Causes, Symptoms,
and Treatment. A Practical Manual for all Fanciers. By Quintin
Craig and James Lyell. In paper, price Is., by post Is. 2d.

POULTRY POR PRIZES AND PROPIT. Contains : Breeding
Poultry for Prizes, Exhibition Poultry and Management of the
Poultry Yard. Handsomely Illustrated. Second Edition. By Prof.
.Tames Long. In cloth gilt, price 2s. 6d., by post 2s. lOd.

RABBIT, BOOK OP THE. A Complete Work on Breeding and
Bearing all Varieties of Fancy Babbits, giving their History, Variations,
Uses, Points, Selection, Mating, Management, &c., &c. SECOND
EDITION. Edited by Kempster W. Knight. Illustrated with
Coloured and other Plates. In cloth gilt, price 10s. 6 d., by post 11s.

RABBITS, DISEASES OP : Their Causes, Symptoms, and Cure.
With a Chapter on The Diseases op Cavies. Beprinted from “ The
Book of the Babbit ” and “ The Guinea Pig for Food, Fur, and Fancy.”
In paper, price Is ., by post Is. 2d.

RABBIT-PARMING, PROFITABLE. A Practical Manual, show¬
ing how Hutch Babbit-farming in the Open can be made to Pay Well.
By Major G. F. Morant. In paper, price Is., by post Is. 2d.

RABBITS POR PRIZES AND PROFIT. The Proper Manage¬
ment of Fancy Babbits in Health and Disease, for Pets or the Market,
and Descriptions of every known Variety, with Instructions for Breed¬
ing Good Specimens. Illustrated. By Charles Batson. In cloth
gilt, price 2s. 6d., by post 2s. 9d. Also in Sections, as follows:—

General Management of Rabbits. Including Hutches, Breed¬
ing, Feeding, Diseases and their Treatment, Babbit Courts, &c. Fully
Illustrated. In paper, price Is., by post Is. 2d.

Exhibition Rabbits. Being descriptions of all Varieties of
Fancy Babbits, their Points of Excellence, and how to obtain them.
Illustrated. In paper, price Is., by post Is. 2d.

REPOUSSE WORK POR AMATEURS: Being the.Art of Orna-
menting Thin Metal with Baised Figures. By L. L. Haslope. Illus¬
trated. In cloth gilt, price 2s. 6d., by post 2s. 9d.

ROSES FOR AMATEURS. A Practical Guide to the Selection and
Cultivation of the best Boses. Illustrated. By the Bev. J. Honywood
D’Ombrain, Hon. Sec. of the Nat. Bose Soc. In paper, price Is., by
post Is. 2d.

SAILING GUIDE TO THE SOLENT AND POOLE

HABBOUB, with Practical Hints as to Living and Cooking on, and

170, Strand, London, W.C.

Working a Small Yacht. By Lieut.-Colonel T. G. Cuthell.
Illustrated with Coloured Charts. In cloth, price 2s. 6 d., by post 2s. 9 d.

SAILING- TOURS. The Yachtman’s Guide to the Cruising Waters
of the English and Adjacent Coasts. With Descriptions of every Creek,
Harbour, and Roadstead on the Course. With numerous Charts printed
in Colours, showing Deep water, Shoals, and Sands exposed at low water,
with sounding. In Crown 8vo., cloth gilt. By Frank Cowper, B.A.

Vol. I., the Coasts of Essex and Suffolk, from the Thames to Aid-
borough. Six Coloured Charts. Price 5s., by post 5s. 3d.

Vol. II. The South Coast, from the Thames to the Seilly Islands, with
twenty-five Charts printed in Colours. Price 7s. 6d., by post 7s. lOd.

Vol. III. The Coast of Brittany: Descriptions of every Creek,
Harbour, and Roadstead from L’Abervrach to St. Nazaire, and an
Account of the Loire. With twelve Charts, printed in Colours. Price
7s. 6d., by post 7s. lOd.

Vol. IV. The West Coast, from Land’s End to Mull of Galloway,
including the East Coast of Ireland. With thirty Charts, printed
in Colours. Price 10s. 6d., by post 10s. lOd.

ST. BERNARD, THE. Its History, Points, Breeding, and Rearing.
By Hugh Dalziel. Illustrated with Coloured Frontispiece and Plates.
In cloth, price 2s. 6d., by post 2s. 9d.

ST. BERNARD STUD BOOK. Edited by Hugh Dalziel. Price
3s. 6d. each., by post 3s. 9d. each.

Vol. I. Pedigrees of 1278 of the best known Dogs, traced to their
most remote known ancestors, Show Record, &c.

Vol. II. Pedigrees of 564 Dogs, Show Record, &e.

SEA-PISHING POR AMATEURS. Practical Instructions to
Visitors at Seaside Places for Catching Sea-Fish from Pier-heads, Shore,
or Boats, principally by means of Hand Lines, with a very useful List of
Fishing Stations, the Fish to be caught there, and the Best Seasons.
By Frank Hudson. Hlustrated. In paper, price Is., by post Is. 2d.

SEA-PISHING ON THE ENGLISH COAST. The Art of Making
and Using Sea-Tackle, with a full account of the methods in vogue
during each month of the year, and a Detailed Guide for Sea-Fishermen
to all the most Popular Watering Places on the English Coast. By
F. G. Aelalo. Illustrated. In cloth, price 2s. 6 d., by post 2s. 9 d.

SEASIDE WATERING PLACES. A Description of the Holiday
Resorts on the Coasts of England and Wales, the Channel Islands,
and the Isle of Man, giving full particulars of them and their attrac¬
tions, and all information likely to assist persons in selecting places in
which to spend their Holidays according to their individual tastes.
Illustrated. Seventh Edition. In cloth, price 2s. 6d., by post 2s. lOd.

SHADOW ENTERTAINMENTS, and How to Work Them: being
Something about Shadows, and the way to make them Profitable and
Funny. By A. Patterson, In paper, price Is., by post Is. 2d.

SHAVE, AN EASY : The Mysteries, Secrets, and Whole Art of, laid
bare for Is., by post Is. 2d. Edited by Joseph Morton.

SHEET METAL, WORKING IN: Being Practical Instructions for
Making and Mending Small Articles in Tin, Copper, Iron, Zinc, and
Brass. Illustrated. Third Edition. By the Rev. J. Lukin, B.A. In
paper, price Is., by post Is. Id.

SHORTHAND, ON GURNEY’S SYSTEM (IMPROVED),

LESSONS IN: Being Instructions ip the Aft of Shorthand Writing as

Published by L. Upcott Gill,

used in the Service of the two Houses of Parliament. By It. E. Miller.
In paper, price Is., by post Is. 2d.

SHORTHAND, EXERCISES IN, for Daily Half Hours, on a Newly-
devised and Simple Method, free from the Labour of Learning. Illus¬
trated. Being Part II. of “ Lessons in Shorthand on Gurney’s System
(Improved).” By R. E. Miller. In paper, price 9 d., by post 10 d.

SHORTHAND SYSTEMS; WHICH IS THE BEST? Being a
Discussion, by various Experts, on the Merits and Demerits of all the
principal Systems, with Illustrative Examples. Edited by Thomas
Anderson. In paper, price Is., by post Is. 2d.

SKATING CARDS: An Easy Method of Learning Figure Skating, as
the Cards can be used on the Ice. In cloth case, 2s. 6 d., by post 2s. 9d.;
leather, 3s. 6cL, by post 3s. 9d. A cheap form is issued printed on paper
and made up as a small book, Is., by post Is. Id.

SLEIGHT OF HAND. A Practical Manual of Legerdemain for
Amateurs and Others. New Edition, Revised and Enlarged. Profusely
Illustrated. By E. Sachs. In cloth gilt, price 6s. 6 d., by post 6s. lOd.

SNAKES, MARSUPIALS, AND EIRDS. A Charming Book of
Anecdotes, Adventures, and Zoological Notes relating to Snakes, Marsu¬
pials, and Birds. A capital Book for Boys, and all interested in Popular
Natural History. By Arthur Nicols, F.G.S., F.R.G.S., &c. Hlus-
trated. In cloth gilt, price 3s. 6 d., by post 3s. lOd.

TAXIDERMY, PRACTICAL. A Manual of Instruction to the
Amateur in Collecting, Preserving, and Setting-up Natural History
Specimens of all kinds. With Examples and Working Diagrams. By
Montagu Browne, F.Z.S., Curator of Leicester Museum. Second
Edition. In cloth gilt, price 7s. 6 d., by post 7s. lOd.

THAMES GUIDE BOOK. From Lechlade to Richmond. For Boating
Men, Anglers, Picnic Parties, and all Pleasure-seekers on the River.
Arranged on an entirely new plan. Second Edition, profusely illustrated.
In paper, price Is. by post Is. 3d.; cloth, Is. 6 d., by post Is. 9d.

TOMATO AND PRUIT GROWING as an Industry for Women.
Lectures given at the Forestry Exhibition, Earl’s Court, during July
and August, 1893. By Grace Harriman, Practical Fruit Grower
and County Council Lecturer. In paper, price Is., by post Is. Id.

TOMATO CULTURE POR AMATEURS. A Practical and very
Complete Manual on the Subject. By B. C. Ravenscroft. Illustrated.
In paper, price Is., by post Is. 3d.

TRAPPING, PRACTICAL: Being some Papers on Traps and
Trapping for Vermin, with a Chapter on General Bird Trapping and
Snaring. By W. Carnegie. In paper, price Is., by post Is. 2d.

TURKEY, THE. A Handy Manual for both the Amateur and Pro¬
fessional Breeder of the Turkey, describing its Characteristics, Varieties,
and Management. By W. WilliS-Harris, Vice-President of the
Turkey Club. In paper, price Is., by post Is. 2d.

TURNING FOR AMATEURS : Being Descriptions of the Lathe and
its Attachments and Tools, with Minute Instructions for their Effective
Use on Wood, Metal, Ivory, and other Materials. Second Edition,
Revised and Enlarged. By James Lukin, B.A. Hlustrated with 144
Engravings. In cloth gilt, price 2s. 6d., by post 2s. 9d.

TURNING LATHES. A Manual for Technical Schools and Apprentices.
A guide to Turning, Screw-cutting, Metal-spinning, &c. Edited by
James Lukin, B.A. Third Edition. With 194 Illustrations. In cloth
gilt, price 3s., by post 3s. 3d.

170, Strand, London, W.C,

VAMP, HOW TO. A Practical Guide to the Accompaniment of Songs
by the Unskilled Musician. With Examples. In paper, price 9d., by
post lOd.

VEGETABLE CULTURE FOR AMATEURS. Containing Concise
Directions for the Cultivation of Vegetables in Small Gardens so as to
insure Good Crops. With Lists of the Best Varieties of each Sort. By
W. J. Mat. Illustrated. In paper, price Is., by post Is. 2 d.

VENTRILOQUISM, PRACTICAL. A thoroughly reliable Guide to
the Art of Voice Throwing and Vocal Mimicry, Vocal Instrumentation,
Ventriloquial Figures, Entertaining, &c. By Robert Ganthony.
Numerous Illustrations. In cloth, price 2s. 6<J., by post 2s. 9 d.

VIOLINS (OLD) AND THEIR MAZERS : Including some
References to those of Modern Times. By James M. Fleming.
Illustrated with Facsimiles of Tickets, Sound-Holes, &c. Reprinted
by Subscription. In cloth, price 6s. 6 d., by post 6s. lOd.

VIOLIN SCHOOL, PRACTICAL, for Home Students. Instructions
and Exercises in Violin Playing, for the use of Amateurs, Self¬
learners, Teachers, and others. With a supplement on “ Easy Legato
Studies for the Violin.” By J. M. Fleming. Demy 4fo, price 9s. 6d.,
by post 10s. 4d. Without Supplement, price 7s. 6d., by post 8s. Id.

WAR MEDALS AND DECORATIONS, A Manual for Collectors,
with some account of Civil Rewards for Valour. Beautifully Hlustrated.
By D. Hastings Irwin. In cloth, price 7s. 6 d., by post 7s. lOd.

WHIPPET AND RACE-DOG, THE : How to Breed, Rear, Train,
Race, and Exhibit the Whippet, the Management of Race Meetings,
and Original Plans of Courses. By Freeman Lloyd. In cloth gilt,
price 3s. 6d., by post 3s. lOd.

WILDFOWLING, PRACTICAL : A Book on Wildfowl and Wildfowl
Shooting. By Hy. Sharp. This work is the result of 25 years’ ex¬
perience in Wildfowl Shooting under all sorts of conditions of locality
as well as circumstances, and can therefore be relied on as a safe and
practical guide. The text is elucidated throughout in the most complete
manner, both as to the birds themselves and the method of approaching
them, the guns and tackle used, and all other points of practical interest.
In short, this book will be the most complete and practical since the
days of Col. Hawker. Demy 8vo, price 12 s. 6 d., by post 13s.

WINDOW TICKET WRITING. Containing full Instructions on the
Method of Mixing and Using the Various Inks, &c., required, Hints on
Stencilling as applied to Ticket Writing, together with Lessons on Glass
Writing, Japanning on Tin, &c. Especially written for the use of
Learners and Shop Assistants. By Wm. C. Scott. In paper, price Is.,
by post Is. 2 d.

WIRE AND SHEET GAUGES OF THE WORLD. Compared
and Compiled by C. A. B. Pfeilschmidt, of Sheffield. In paper,
price Is., by post Is. Id.

WOOD CARVING FOR AMATEURS. Full Instructions for pro¬
ducing all the different varieties of Carvings. 2nd Edition. Edited by
D. Denning. Price Is., by post Is. 2d.

PUBLISHED BY E. and F. N. SPON

JtECENTL 1 P UBLISHED.

Crown 4to, full gilt, fancy cloth, 478 pages Letterpress and 735 Engravings, price 7 / 6 .

POPULAR ENGINEERING:

being interesting and instructive examples in

CIVIL, MECHANICAL, ELECTRICAL, CHEMICAL, MINING,
MILITARY, and NAVAL ENGINEERING,

GRAPHICALLY AND PLAINLY DESCRIBED AND

Specially Written for those about to enter the Engineering Profession and the
Scientific Amateur. With Chapters upon

PERPETUAL MOTION and ENGINEERING COLLEGES and SCHOOLS.

By nr". DYE.

MECH ANICS’ OWN BOOK.

SPONS’ MECHANICS^WN BOOK:

A MANUAL FOR HANDICRAFTSMEN AND AMATEURS.

Complete in One large Vol., demy 8vo, cloth, containing 700 pp. and 1420 Illustrations.
Fourth Edition, 6/-; or half-bound, French morocco, 7/6,

Contents:

Mechanical Drawing; Casting and Founding in Iron, Brass, Bronze, and other Alloys;
Forging and Finishing Iron; Sheet Metal Working; Soldering, Brazing, and Burning;
Carpentry and Joinery, embracing descriptions of some 400 woods; over 200 Illustrations of
Tools and their TJ?es; Explanations (with Diagrams) of 116 Joints and Hinges, and Details of
Construction of Workshop Appliance;; Bough Furniture, Garden and Yard Erections, and
House-Building; Cabinet-making andVeneering; Calving and Fret-cutting; Upholstery;
Painting, Graining and Marbling; Staining Furniture, Woods, Floors and Fittings ; Gilding,
Dead and Bright, on various Grounds; Polishing Marble, Metals and Wood; Tarnishing;
Mechamcai Movements, lUustrating contrivances for transmitting Motion; Turning in Wood
with Thatch ^'“^“^Tereaj-ootta^ and Concrete ^Roofing

Plastering and -’ - ’ ”

Electric Sys

Ditches and_

New Countries.

HOUSEHOLD MANUAL.

SPONS’ HOUSEHOLD MANUAL:

A TREASURY OF DOMESTIC RECEIPTS AND GUIDE FOR HOME
MANAGEMENT.

Demy 8vo, cloth, containing g75 pp. and 250 Illustrations, price 7/6; or half-bound,
French moroeeo, 9 -

Principal Contents:

. Hints f ° r selecting a good House; Sanitation; Water Supply; Ventilation and Warming ;
F 'Shfang; Furniture and Deem-ation; Thieves and Fire; The Larder; Curing Foods for
lengthened Preservation; The Dairy; The Cellar; The Pantry; The Kitchen; Receipts for
TOom e - S: The e ^d“n e o^ e Tte°lSrsSy“ S ^he P SGS ““*?*&£* The Dming;room ; T The Drawing-
^ayground; The Work-roi
sehold Law.

London: E. & F. N. SPON, 125, Strand.

New York: SPON & CHAMBERLAIN, 12, Cortlandt Street.

ROWLANDS’ ARTICLES

For the HAIR, COMPLEXION, & TEETH, are the

PUREST AND BEST.

Rowlands’ odcnto,

An antiseptic, preservative, and aromatic den¬
tifrice, which whitens the teeth, prevents and
arrests decay, and sweetens the breath. It
contains no mineral acids, no gritty matter or
injurious astringents, keeps the mouth, gums,
and teeth free from the unhealthy action of
germs in organic matter between the teeth,
and is the most wholesome tooth-powder for
smokers. It is most beautifully perfumed,
and is a perfect luxury for the toilet-table of
everybody. 2s. 9d. per box.

ROWLANDS’ MACASSAR OIL

Is the best preserver and beautifier of the hair of children and adults ; prevents
it falling off or turning grey, eradicates scurf and dandruff, and is also the best
brilliantine for ladies’ and everybody’s use, and as a little goes a very long way
it really is most economical for general use ; is also sold in a golden colour for
fair-haired ladies and children ; it contains no lead or mineral ingredients, has
a most delightfully fragrant bouquet of roses, and is considered the most perfect
toilet luxury ever produced. Bottles, 3s. 6d., 7s., 10s. 6d.

ROWLANDS’ KALYDOR,

A most cooling, soothing, healing, and refreshing preparation for the Skin and
Complexion of Ladies, and all exposed to the summer sun and dust, or the cold
and damp of winter; it is warranted free from all mineral or metallic ingre¬
dients, or oxide of zinc, of which most Cosmetics are composed, and which ruin
the skin. It effectually disperses Chaps, Chilblains, Freckles, Tan, Sunburn,
Stings of Insects, Redness, Roughness of the Skin; relieves Irritation of the
Skin, Prickly Heat, &c., renders the

SKIN SOFT AND SMOOTH,
and produces a beautiful, pure, and delicate complexion. Size 4s. 6d. and
8 s. 6d. ; half-sized bottles, 2s. 3d.

ROWLANDS’ ESSENCE OF TYRE

effectually dyes red or grey hair a permanent brown or black. 4s.

ROWLANDS’ EUKON1A.

A pure Toilet Powder in three tints, White Rose, and Cream, for ladies of a
Brunette complexion and those who do not like white powder. Boxes, Is.;
large boxes, 2s. 6d. _

Ask Chemists for ROWLANDS’ ARTICLES, of 20, HATTON
GARDEN, LONDON, and avoid spurious imitations.

SPRATTS PATENT LIMITED, BERMONDSEY, S.E.

Pamphlet on CANINE DISEASES GRATIS.

!l>

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