Insect Physiology (Wigglesworth)

INSECT PHYSIOLOGY
BY
V. B. WIGGLESWORTH, M.A., M.D.
LEaruRIilB. AIID IIILIlIilR RIilSEARCH PBLIoOW III MEDICAL liill!rOJIOLOOY.
LONDOII SOIlOOL 01' HYGIRIIR AND TROPIOAL IIRDICIN Iil
WITH 13 ILLUSTRATIONS
METHUEN & CO. LTD.

36 ESSEX STREET W.C.
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To the Insect, order and disorder are exposed to sight;
Bnd so we think to see the little emmets confer
and locking their antennae ImmediatIy transmit
the instInctIv call. which each and all CBIl feel; where••
the mutual fellowship of distributed cells
hath so confounded thought that explanation Is Cetch'd
from chemic agency: because ill that science
the read-Ion of unknowII fol'C8l! Is described and 8umm'd
In mathematic fonnulae prellnant of truth,
and of such universal scope that, being caU'd laws,
their mere description passeth for Efficient Ca.use.
-1l0BERT BRIDGES
INTRODUCTION
T HE fundamental ·processes of vital activity,

·the ordered series of physical a~d chemical
changes which liberate energy anp_ maintain the
'immanent movement' of life, are probably the
same wherever 'living matter' exists. The
description of these changes is the ultimate goal
of physiology, of whatever group of organisms;
but the tissues of insects hJtve been so little
studied from that point of view, that in this
book we shall be concerned only with physiology
on a humbler plane: with the grosser fUD,ctions
of the organs and tissues, and with the mecha-
nisms by which these functions are co-ordinated
to serve the purpose of the insect as a whole.
Of all the zoological classes, the insects are the
most numerous in species and the most varied
in structure; the general physiology of the group
is therefore only too apt to be obscured by the
endless specializations of particular forms; and
v
vi INSECT PHYSIOLOGY
the main difficulty of a work like the present is
the exclusion of all that is special and non-
essential, and the retention of only that material
which best illustrates the general theme. There
are, indeed, certain common factors which con-
dition the physiological make-up of the insects,
and these factors serve, to some extent, to liIik
them all together into one system. They are
essentially terrestrial animals. This circumstance
determines the characters of their cuticle, and
this, in tum, conditions their respiratory mecha-
nist;ll and the physiology of their growth. The
respiratory mechanism and the cuticular skeleton
are among the factors which restrict the size of
insects. Their small size and terrestrial habit
render them very prone to lose water; and the
urgent need for the conservation of water in-
fluences the respiratory, excretory and digestive
systems. All these systems show special changes
when the insect reverts to an aquatic life.
Yet, although the interaction of these factors
results in some uniformity of principle in the
physiology of insects, their diversity in habit,
food and environment causes such endless varia-
tion in detail, that, quite apart from the con-
spicuous gaps in our present knowledge, any
limited treatment of the subject must in any case
be more or less arbitrary in presentation; and,
were there space to insert them, nearly all the
generalizations that are attempted should have
qualifying instances.
_The history of insect physiology is peculiar.
The early microscopists, Hooke, Malpighi, Leeu-
wenhoek, made many observations on the
st.ructure of insects. and many accurate inferences
IN'rB.ODUC'rION vii
about their physiology. More was added by the
great naturalists, Swammerdam, Reaumur, de
Geer. But, with such conspicuous exceptions as
Newport, Graber, Lubbock, Plateau, and others
of more modern times, the majority of entomo--
logists, until recent years, have been so fully
occupied with the morphology and taxonomy of
their colossal group that such advances in phy-
siology as have been made have commonly been
mere by-products of morphological study. From
time to time we find the physiologists of the last
century, Dutrochet, Treviranus, Marshall Hall,
v. Kolliker, Claude Bernard, turning to the
insects to illuminate their theme; but their
concern was not with the insect as such.
Within recent years, interest in the physiology
of insects has arisen in a new quarter. The applied
entomologist, confronted with the ravages of in-
sects in the spheres of agriculture and of public
health, has wanted to know something about their
nutrition, about the laws governing their responses
to sensory stimuli, about their reactions to
parasites, about the precise way in which their
bodies are adapted to diverse climatic conditions,
and about the action upon them of toxic sprays
and gases. With this demand for increased know-
ledge has come a realization of our present
ignorance.
The sketch which follows is based on the study
of nearly 2,000 publications, and on a certain
amount of original work. It has not been possible
to include references to all these sources; some
300, only, have been selected-most of them
recent works which will introduce the student to
the earlier literature. For this reason, reference
viii INSECT PHYSIOLOGY
to a given author is not intended to imply that
this author was necessarily the first to have made
the observations in question; many such refer·
ences are to textbooks and reviews.
CONTENTS
0iW'. PAGE
IN'l'RODUCTION V
I Tro: INTEGUMENT 1
Composition of the Cuticle. Structure of
the Cuticle. Surface Forces and the Insect
Cuticle. The Moulting, Formation and Pig-
mentation of the Cuticle. The Mechanism
of Moulting and Hatching. Other Dermal
Glands.
IT RESPIRATION 16
The Tracheae and Tracheoles. The Dif-
fusion Theory of Insect Respiration. The
Function of the Spiracles. Ventilation o£..the
Tracheal System. The Regulation of Res-
piration. Respiration of Aquatic Insects.
Respiration of Parasites.
III THE CIRCULATORY SYSTEM .AND THE BLOOD 32

Mechanism of the Circulation. Control of
the Heart-beat. Composition and Properties
of the Blood. Cytology of the Blood. Func-
tiOIis of the Blood.
IV DIGESTION 42
The Alimentary Canal. Proventrioulus.
Peritrophio Membrane. Salivary Glands.
Secretion. Reaction of the Gut. Digestive
Enzymes. Symbionts in Digestion. Absorp-
tion.
V EXCRETION 56
Excretion of Dyes. Nephrocytes. Urate
Cells. Pigments. Verson's Glands. Mal-
pighian Tubes and the Urine. Excretion in
ix
x INSECT PHYSIOLOGY
ellAP. rAom
Rhodniu8. Excretion in other Insects. Ex-
cretion in Aquatic Insects. Accessory Func-
tions of the Malpighian Tubes.
VI NUTRITION AND METABOLISM. 70

Food Requirements. Vitamirra and Sym-
biotic Organisms. Water Metabolism.
Temperature of Insects. Respiratory Meta-
bolism. Anaerobic Metabolism. Chemical
Changes during Metamorphosis. Suspended
Metabolism. Pigment Metabolism.
VII REPRODUCTION AND GROWTH • 86

Growth. Moulting. Metamorphosis. His-
tolysis and Histogenesis. Regeneration.
Determination of Sex. Determination of
other Characters.
VIII THE NERVOUS SYSTEM, SENSE ORGANS AND

BEHAVIOUR 100
The Nerve Cord. The Brain. Reftex
responses in the Normal Insect. Orientation.
Sense Organs : Mechanical Senscs. Chemical
Senses. Vision. Colour Vision. Insect
Behaviour.
REFERENCES 116
INDEX 130
INSECT PHYSIOLOGY
CHAPTER I
THE INTEGUMENT
T HE key to much of the physiology of insects is

to be found in the nature of their cuticle. As
was first shown by Haeckel, the cuticle consists of
non-living material, the product of a living matrix
-a single layer of epidermal cells. Its most familiar
constituent is the nitrogenous polysaccharide---chitin ;
but neither qualitatively nor, in some cases, even
quantitatively, can chitin be considered its most
important element. For chitin rarely makes up more
than 50 per cent. of the cutic'le; more often it forms
only about 30 per cent. 53, 152; and the most char-
acteristic properties of the cuticle, its rigidity and its
impermeability, are not dependent on the chitin
present. For those parts which are richest in chitin,
the inner layers, are the most flexible and elastic;
and where chitin is present without the other con-
stituents, as in the anal papillae of mosquito larvae
(p. 68) or in the peritrophic membrane of the in-
testine (p. 46), the structure is freely permeable to
water and to many substances in solution.
Composition of the Cuticle. The cuticle is in-
deed a very complex tissue. It consists of two
primary layers: a relatively thick endocuticle, known
also as the 'Hauptlage' or 'Innenlage', composed
of chitin and of an insoluble protein; and a very
thin epicuticle or 'Grenzlamelle' 188. This latter
1
2 INSECT PHYSIOLOGY
contains neither protein nor chitin; it is only about
1,1.1. in thickness; yet it appears to confer upon the
insect cuticle its essential property of impermeability.
For the substance of this epicuticle the name e cuti-
culin.' has been proposed 288. It bears much the
same relation to the underlying chitin layer as the
delicate cuticle of the plant does to cellulose. Chemi-
cally, also, it seems closely allied to the plant cuticle,
resisting solution in ooncentrated sulphuric or hydro-
chloric acids, in which the endocuticle dissolves, but
dissolving in caustic alkalies, to which chitin is resis-
tant. It is not a protein nor a carbohydrate nor a
simple fat; perhaps it is a complex mixture of
combined fats or waxes 188. It is evident that the
biological importance of this remarkable substance
makes it a most desirable subjeot for chemical study.
The protein of the endocuticle is readily demon-
strated by the protein colour tests: the biuret,
Millon's and xanthoproteic reactions, all of which
are intensely positive 288. The best test for chitin
is that of van Wisselingh, which consists in its con-
version into chitosan by strong caustio potash at a
high temperature, and the recognition of chitosan
by its solubility in acids and by the violet coloUl it
gives with iodine 0.3. The cuticulin of the epiouticle
is a less well-defined substance; quite likely it is a
mixture; its most characteristic properties are its
insolubility in strong mineral acids, and its rapid
fusion into oily droplets when heated in the presence
of caustic potash and potassium chlorate (cerinic acid
test) 188.
In unstained sections, the endocuticle is colourless,
the epicuticle typically amber-coloured. But in
many parts, where the cuticle is rigid and inflexible
or e sclerotized " the outer part of the endocuticle,
also, is impregnated with the amber-coloured cuti-
culin, often more or less mixed with melanin; and
thus a third layer is formed in which chitin, protein,
e cuticulin ' and other substances are present. This
THE INTEGUMENT 3
layer is responsible for the rigidity of the cuticle ;

it is variously termed the exoouticle, 'Emailschicht '
or 'Lackschicht' 286.
In a few insects the cuticle contains lime. Some·
times this forms a superficial deposit on the spines·
of aquatic larvae (Pericoma, Psychodidae, Dipt.)
from waters rich in calcium bicarbonate 166; some·
FIG. I.-The insect cuticle: semischematic

br, hair or bristle; d, duct of dermal gland; end, endocuticle; BP, epi-
cuticle; ez, exocuticle; VI, dermal gland; hllP, epidermis (hypodermis) ;
pc, pore canals; ,ce, .. Sekretschlcht "; 10, tormogen or socket-forming
Cell; Ir, trichogen or hair-forming ceIl.
times it is laid down, before moulting, in pits in the

new cuticle called calcareous warts (Stratiomyidae,
Dipt.) 198. •
Finally, there is sometimes a layer whioh is believed
to be formed by the outpouring of secretion from the
dermal glands at the time of moulting. This is
termed the' Sekretschicht '. It is not always present,
and sometimes it amounts to nothing more than a.
4 INSECT PHYSIOLOGY
little dried secretion of no functional signi1icance 288 ;
but sometimes it is an important layer; and in the
case of many of the brilliantly coloured Cicindelidae,
it is ·apparently the layer which is responsible for the
metallic tints 24D. Its chemical nature is unknown.
Structure of the Cuticle. The histological
structure of the cuticle is no less complex than its
chemistry. It is most commonly described from
stained sections, and in these the epicuticle and exo-
cuticle, which do not stain, appear homogeneous and
structureless; the endocuticle shows more or less con-
spicuous horizontal lamellae. In some preparations
the endocuticle shows also very faint vertical stria-
tions-the so-called ' pore-canals'. The older histo-
logists paid considerable attention to these structures;
but in sections prepared by modern histological
methods they are so inconspicuous that most recent
authors have neglected them, regarding them simply
as processes (pseudopores) from the epidermal cells,
which have become chitinized. But if the cuticle
of certain insects (Rhodniu8, Hem. 286, Tenebrio larva,
Co]. 288, Periplaneta, Orth. 288) is cut in the fresh state
with the freeZing microtome and mounted without
staining, the pore-canals are very conspicuous objects
(often far more conspicuous than the horizontal
lamellae) running right through the endo- and exo-
cuticle, though not piercing the epicuticle (Fig. 1).
Moreover, the canals are patent, and contain proto-
plasmic processes or fluid; for if the sections are
allowed to dry in the air before being mounted, the
canals become filled with fine threads of gas. The
cuticle is therefore not entirely dead; like the bone
and dentine of vertebrates, it probably contains
cytoplasmic filaments. .
In the past there has been a. certain amount of
controversy as to whether, microscopically, the sub-
stance of the endocuticle has a foam-like structure
or a fibrillar structure 168. At the present time one
would probably be safe in dismissing both these views
THE tNTEQt1MENT 5
a.nd regarding this substance as an elastic gel, which
in many insects is formed around protoplasmic fila-
ments from the epidermal cells (the pore-canals), and
in nearly all cases is, by some unknown mechanism,
laid down in a succession Qf lamellae. The extent
to whjch the endoc,"!lticle can be stretched, when it
is not impregnated With cuticulin, is often astonish-
ing: the first stage larva of ,Rhodniu8 can receive into
its abdomen more than twelve times its own weight
of blood. The epicuticle, on the other hand, appar-
ently cannot be stretched; it is often highly con-
voluted, and it accommodates itself to changes in
volume by the smoothing out of the folds in its
surface 286.
A rather special arrangement of the endocuticle is
met with in the rigid armour of beetles. Here each
layer of the endocuticle is made up of a series of
parallel rodlets (' Balkenlagen ') with interstitial sub-
stance of a different nature between them. The
'Balken' of each successive layer are set at an angle
to those above and below; an arrangement which
probably increases the inflexibility of the cuticle. In
these cases the pore-canals run upwards between the
criss-crossing.' Balken " appearing in surface view as
stellate points 240, 23!,.
Surface Forces and the Insect Cuticle. Since
the surface of a body varies with the square of its
l'adius, whereas its volume or mass varies with the
cube of its radius, it follows that the ratio of surface
to mass becomes progressively greater as the size
of the body diminishes. Consequently, the forces
resident in surfaces become relatively stronger. In
the case of insects, the surface forces are, indeed,
frequently great enough to move the whole mass of
the body; and they introduce a factor into the life
~f insects, partiCUlarly when they come into contact
w{th water, which is unnoticed by larger animals.
It is necessary, therefore, to consider the properties
of the cuticle from this point of view.
6 INSECT PHYSIOLOGY
As a general rule, the cuticle is not readily wett~d'
. by water; there is, in fact, a large' angle of contac~ ,
between the water surface and the cuticle (Fig. 2,
A) 37. Consequently, when an insect stands on
water, the surface tension acts in the opposite direc-
tion to gravity, and the insect is held up as though
by an elastic membrane. This' hydrophobe' pro-
perty may belong to the epicuticle; but in some cases•.
at least it is accentuated by the presence of glands.
Fig. 2.-Diagrams illustrating the properties of the inseot

cuticle in oontact with w:ater I
A, hydrophobe object supported by the surface tension of water ; B, resplra·
tory siphon with hydrophobe rim similarly supported; C, surface forces
acting on a hydrophobe heir brought obliquely to the water surface; D, a
crown of such heirs supporting a respiratory siphon; E, hydrophobe hairs
brought vertically against the water surface; F, Insect body with a ventral.,
covering of such heirs carrying a layer of air. .
produoing a fatty secretion 2'. That is the case, for

example, around the spiracles of Muscid larvae, and
around the respiratory tube of the larva of Eri8taZiB,
by means ·of which this insect, like ma.ny oth~r
aquatic larvae, is enabled to hang suspended fro'"
the surface film with the remainder of its body sul
merged (Fig. 2, B) 282. ..
The effect of surface forces becomes still mOI;8"
evident when hairs or bristles are present on the bod;(
THE INTEGUMENT 7
Where a fine pile of these hyarophobe hairs is set
vertically on the body surface, they hold the water
t a distance, so that the insect, when it goes below
Ghe water surface, can carry with it a film of air
(Fig. 2, E, F). We shall see that this film has some
·emarkable properties in relation to the respiration
If aquatic forms (p. 29). Then, if a single bristle
inclined at an angle-for simplicity we will suppose
~ an angle of 45°, and we will suppose that the angle
of contact is also 45°-and brought up to the water
surface from below, as can be seen from Fig. 2, C,
Lhe surface tension of the water will draw it over so
."to increase the angle of inclination. It will behave
as though tho one surface were hydrophile and the
ther hydrophobe. Many insects hold themselves at
be water surface for purposes of respiration by means
~ rows or circles of such bristles (Fig. 2, D) 37.
l'errestrial insects avail· themselves of surface
:-.ces ill' quite a different way: namely, in order to
~ to surfaces too smooth to provide a purchase
their claws. 'l'he physical mechanism of the
..LIlS (pulvilli and such like) that are used for this
lrpose is by no meallil certain in all cases; but in
ae example that has been recently studied (that of
e bug Rhodnius) 289, it has been shown that the
.3ect is held by the adhesion or seizure of the climbing
'''gan to the surface. In other words, it is the surface
llecular forces that are directly concerned in sup-
l-ting the body weight.
The Moulting, Formation and Pigmentation
of the Cuticle. Through the possession of a more
')r iess rigid carapace, incapable of increasing its
surface area once it has been laid down, the insect
curs the necessity for discontinuous growth. In
1 pages (p. 89) ·we shall consider some of the phy-
~ical processes which lie at the back of insect
"Gh; but one change which it always involves
,~ casting of the old cuticle, and the formation
a" new and larger one. This side of growth, or
"
8 INSECT PHYSIOLOGY
ecdysis, may be conveniently considered here, for
it throws a good deal of light upon the properties of
the insect cuticle.
Chitin is a nitrogenous substance, though its
nitrogen content is relatively small (about 8 per cent.
-only half that of protein), and the suggestion has
therefore frequently been made that it is really a
waste product, and moulting or ecdysis an excretory
phenomenon. But this idea fails to take into account
the fact, which has long been known, that during
moulting most of the old cuticle is dissolved, and is
not, in fact, lost to the insect. In a recent study of
mGulting in the bug Rhodniu8, it has been shown that
only about 14 per cent. of the cuticle of the abdomen
is actually shed; the remaining 86 per cent. is reo
absorbed by the insect and is presumably used again
for the formation of the new cuticle 286.
This process of digestion and absorption of the
cuticle raises many interesting problems in physiology.
(i) Firstly, what is dissolved 1 Only the endocuticle
is dissolved, the chitinous and protein layer; the
epicuticle is quite untouched; and when the outer
part of the endocuticle is impregnated with cuticulin
(that is, when an exocuticle is present) this also is
quite untouched 286. (ii) Se~ondly, how is solution
brought about 1 Between the old cuticle and the
new is a thin layer of fluid, the exuvial or moulting
fluid; this must be responsible for dissolving the
cuticle. Now the moulting fluid is a more or less
neutral fluid incapable of ,~ttacking the chitin and
scleroprotein of the cuticle by simple chemical action;
and it contains no cellular elements. It must there-
fore contain digestive enzymes: a chitinase and a
protease 286. (iii) Thirdly, where do these enzymes
come from 1 Those insects in which the digestion
of the endocuticle is most marked, t.he Lepidoptera 263,
the Coleoptera 211, the Hemiptera 286, possess well-
developed dermal glands whic,p. become active at the
time of moulting; and, at any rate in the case of
THE INTEGUMENT 9
the bug Rhodniu8, the onset of functional activity in
these glands coincides with the commencement of
digestion· of the old cuticle 286. It is therefore highly
probable that they are responsible for the production
of the enzymes in the moulting fluid; although, of
course, this does not exclude the possibility that much
of the fluid itself may be exuded by the general
epidermis 189. (Later (pp. 60, 90) we shall have to
consider other hypotheses as to the function of these
glands.) (iv) Fourthly, by what route are the pro-
ducts of digestion absorbed 1 Now the quantity of
moulting fluid present at any moment is very small ;
it often amounts to little more than a moist film, far
thinner than the old endocuticle that is to be dis-
solved (Fig. 3, A). It is clear, therefore, that the
moulting fluid must be circulating and carrying with
it into the body of the insect the products of digestion.
At the time of moulting in the silkworm, much of this
fluid is swallowed by the mouth 265, but in Rhodnius,
the only other insect in which this problem has
been considered, the absorption seems to take place
through the general surface of the newly-formed
cuticle 286.
This raises the question of the resistance of the new
cuticle to solution by the moulting fluid, its method
of formation, and its permeability,. (i) The new
cuticle resists digestion presumably because the non-
chitinous epicuticle is secreted first; the protein and
chitin of the endocuticle are therefore protected from
the enzymes in the moulting fluid 286. (ii) The method
of formation of the endocuticle probably varies in
different insects, and according to the structures that
are to be formed. Sometimes, as in the developing
bristles, a cellular process gradually becomes impreg-
nated with chitin and cuticulin: these secretions are
laid down intracellularly. Often, the material of the
endocuticle seems to be separated in fluid pr semi·
fluid form around the filamentous processes from the
epidermal cells, which later constitute the 'pore.
10 INSECT PHYSIOLOGY
canals 209.286: it is only a question of words whether
t~is phenomenon is called cellular transformation or
secretion. (iii) The permeability of the cuticle when
newly formed must be very different from its per-
mf e~
end
I
\ ' I
DC
'"
mf
FIG. 3.-A, B, moulting of a cuticle in which an exocuticle
is wanting. {Modified after Wigglesworth.} In A the
new epicuticle is formed; the digestion of old endo·
cuticle has scarcely begun. In B the digestion and
absorption of the old endocuticle is almost complete.
C, D, transverse sections through the prothorax of an
insect at stages corresponding with A and B, showing
the absence of exocuticle at the line of weakness.
(Semischematic.)
end, endocuticle; "P. epicuticle; e"" exocuticle; gl. moulting gland;
mi. moulting fluid; ne, new cuticle; 00, old cuticle; W, line of weakness ill
cuticle.
meability when mature. For we have seen that the

mature epicuticle is impermeable to water; but if
the foregoing account is correct, up to the time when
the old skin is cast, it must remain freely permeable
both to water and to the digestion products of the
THE INTEGUMENT 11
endocuticle. In other words, at the time of moulting
some essential change takes place in the properties
of the epicuticle.
Up to the time of moulting the new cuticle is soft
and almost colourless. Within an hour or two. of the
casting of the old skin, it has both hardened and
darkened; and it is probably during this process that
its impermeability develops. The two processes of
melanin formation and the hardening of the cuticulin
take place at the same time; but they are separate
processes, because the dark areas are not necessarily
harder than the pale 121. Now the formation of
melanin is known to be dependent upon the access
of oxygen; it is natural to suppose that the hardening
of the cuticle is equally so; and the obvious con-
clusion is that the shedding of the old skin exposes
the new skin to the air and both hardening and
blackening can then proceed.
But it is more complicated than that. For if the
new cuticle is exposed to the air by dissecting away
a portion of the old cuticle twenty-four or forty-eight
hours before the insect is due to moult, although the
epicuticle is at this time fully formed, darkening and
hardening do not take place until the insect is ripe
for moulting one or two days later 288. And con-
versely, insects which have failed to moult for various
reasons, may have their new cuticle perfectly ha:r:d-
ened and pigmented beneath the remnants of the old
skin. In other words, the influence which brings
about these changes is exerted from within the insect,
presumably by the epidermal cells. Not uncommonly,
the changes in question are confined to the epicuticle ;
so that the epidermal cells must exert their influence
through the substance of the endocuticle, which is
itself unaffected. (It is true that in many cases the
bulk of the endocuticle is laid down after moulting 238,
and when this is so the epidermal .cells are in direct
contact with the' Pigmentschicht ' during the process
of darkening.) Perhaps we have here the function
of the pore-canals: they enable the epidermal cells
to. effect, from a distance, oxidative enzymic changes
in the epicuticle 88, 286. As we shall see later (p. 82)
it is probably the oxidizing enzyme which is conveyed
by the pore-canals; the substrate or chromogen being
previously localized in the cuticle and so determining
the distribution of the pigmentation 107.
The Mechanism of Moulting and of Hatching.
When the insect has digested its old cuticle and laid
down the new one, it has fresh problems to solve. In
the first place, it has to rupture the old skin and
escape from it. Now any cuticle that has to be shed,
that is, the cuticle in every stage except the adult,
has a 'line of weakness '-sometimes in the head,
most often in the mid-line of the thorax 80. Along
this line the exocuticle is wanting, and the endo-
cuticle extends right up to the very delicate epicuticle
(Fig. 3, C). Consequently, when the endocuticle
is completely dissolved (and the process of moulting
is so timed that this happens just when the new
cuticle is ready) the old skin needs but the slightest
force for it to split along the line of weakness (Fig.
3, D).
The rupturing force is the pressure of the blood.
The insect contracts the abdominal muscles, often
repeatedly in a peristaltic fashion, and so increases the
pressure of blood in the thorax, which bulges out-
wards until the old skin splits. Many insects add to
the efficiency of this process by first swallowing air 38
(as vertebrates fill the lungs with air during defae-
cation or parturition) or, if they are aquatic, by
swallowing water 25. Once the old skin is split, the
insect escapes from it by gentle peristaltic movements,
often aided by gravity-for many insects hang head
downwards during moulting 80.
lt is commonly stated that the moulting fluid
serves as a lubricant to facilitate this process-indeed,
that is often considered to be its chief function. But
in many insects the moulting fluid has practica.lly
THE INTEGUMENT 13
disappeared by this stage; the insect is quite dry,
the inner surface of the old skin only slightly moist 286 ;
and in many aquatic and terrestrial pupae a layer
of air appears before hatching between the pupal skin
and the developed adult-due, no doubt, to the
energetic absorption of the moulting fluid.
When the insect has withdrawn its limbs, and freed
itself completely from the old skin or from the pupa,
it must enlarge the new cuticle rapidly to the required
size, and bring about the expansion of its wings,
before the epicuticle has hardened. This it does in
the same'manner, by swallowing more air or water,
and forcing the body fluid under pressure into the
wings. In the Ephemeroptera and in many Lepi-
doptera the sole function of the gut in the adult
insect is to receive air for this purpose. If the crop
of the cockroach is pricked with a needle, the dis-
tension of the new cuticle cannot take place, and the
insect colJapses like a pricked balloon 80; or if the
tips of the wings in the dragon-fly are cut off, blood
drips from them and they cannot expand 38.
We may note here that the process of hatching
from the egg is similar in many particulars to that of
moulting or emergence from the pupa. It is not
known whether the amniotic fluid in which the embryo
is bathed contains digestive enzymes which weaken
the membranes of the egg, nor whether any of this
fluid is absorbed through the general surface of the
young insect's skin; but in many cases the fluid is
swallowed by the mouth shortly before hatching 290
(like the moulting fluid of the silkworm). Many
insects then tear or bite their way out of the egg.
Others swallow air, and by muscular contraction of
the abdomen, drive the head or thorax against the
shell and cause it to burst open. Certain groups have
special structures, spines, saws, hard plates or dis-
tensible bladders, which enable them to increase the
efficiency of this process by concentrating the impact
at one point of the egg, and so piercing the shell OJ;'
forcing off the egg cap 83, 290. (This last arrangement
finds a parallel in the ptilinum of emergent Muscid
flies, which they distend with blood in order to force
off the cap of the' puparium '80.) When the insect
has left the egg, it commonly swallows air or water
once more, and so increases its bulk before its cuticle
has hardened 290.
Other Dermal Glands. Besides the glands
which digest the old cuticle a.t the time of moulting
(p. 8), and in some cases perhaps produce the
'Se.kretschicht' (p. 3), and those concerned in the
production of hydrophobe substances in the cuticle
(p. 6), there are many other types of dermal glands
in insects. Some of these are very like the moulting
glands in structure; others are more complex multi-
cellular glands; but they are nearly always lined with
a delicate membrane continuous with the epicuticle 269.
These glands have the most varied functions. Some-
times, as in many Coccids, they produce a protective
coating of wax, which may be moulded into beautiful
forms as it passes through the orifices in the cuticle 288.
In the feet of Embioptera and Embiidae (Dipt.) they
are silk glands. Sometimes, as in Lepidoptera, they
are scent glands which occur only in the male sex,
often discharging through specially modified scales
on the wings and elsewhere; they are then generally
credited with an aphrodisiac function 82. Some-
times, as in Hemiptera and Blattidae, they are stink
glands, producing a nauseous secretion which is held
responsible for the immunity from predators which
many such insects enjoy 149. In certain beetles,
notably Chrysomelidae, strongly smelling or bitter
fluids may be ejected by dermal glands provided with
large reservoirs 160. In other insects fluids may be
given out which produce urticarial reactions in the
human skin.
This list by no .means exhausts the multiplicity of
dermal glands already known. But at the present
time, the relation of these glands to the general phy.
THE INTEGUMENT 15
siology of the insect is almost entirely unexplored.
In some cases, as for instance in the larva of M elasoma
(Chrysomelidae, Col.) in which the secretion contains
the aromatic salicin concentrated from the willow
leaves on which it feeds 259, it appears as though the
insect has found a biological use for the waste products
of its metabolism; but it cannot at present be
claimed that that is so with the wax or other types
of dermal glands.
CHAPTER II
RESPffiATION
ERRESTRIAL life is a perpetual conflict be-
T._ tween the need for oxygen and the need for
water; for those conditions which favour the entry
of oxygen into the body, favour also the escape of
water_ This conception, applied to the insects, illu-
minates the whole subject of their respiration 115.
The majority of insects, as we have seen, have become
encased in a waterproof covering; a covering which
is relatively imperm.eable to oxygen also. They
breathe, as was discovered by Malpighi, through
tracheal tubes, which open along the sides of the body
through a series of spiracles, and convey air directly
to the tissues (Fig. 5, A). The tracheal trunks anas-
tomose one with another, and then branch and
rebranch, becoming finer and finer, until their ultimate
ramifications either penetrate the individual fibres of
the muscles or the cells of other tissues, or form a
rich network over their surface 282.
The Tracheae and Tracheoles. Morphologi-
cally, the tracheae are invaginations of the ectoderm.
Consequently, they consist of a cellular matrix and
a lini~ cuticle_ As the result, no doubt, of the
physical stresses to which it is subjected during its
formation, the cuticle is thrown into spiral folds, and
sometimes, probably because the cells have continued
to secrete the cuticular substance after the folds have
developed, the final result appears as though the
tracheae were made up of a uniform membrane,
thickened with spiral filaments or taenidia 282. This
16
RESPIRATION 17
spiral formation enables the tracheal tubes to be
bent and pressed upon without collapsing.
The. chemistry of the tracheal outicle resembles ·that
of the general surface of the insect. The innermost
layer is composed of the non-chitinous' cuticulin ' ;
in the larger trunks a certain amount of chitin is
present, presumably deeper to this 152; but in the
small branches chitin is absent 53. Whether the sub-
stance of the finest branches is protein, or is related
to 'cuticulin', is not known. .
When the tracheal branches become reduced to
about l,u in thickness, they no longer show any spiral
folding, and are then termed tracheoles or tracheal
capillaries. Sometimes the transition from trachea
to tracheole takes place gradually; but sometimes
a tracheal branch of 2-5,u in diameter, after entering
a large stellate 't_raoheal cell', breaks up abz:uptly
into a sheaf of fine tracheoles. On account of the
readiness with which this tracheal cell stains with
osmic acid, special.properties have been attributed
to it in connexion with respiration and tissue oxida-
tions. But the evidence for these hypotheses is very
slight; the tracheal cells are often absent; and
altogether it is more probable that they are merely
matrix cells of tracheae or tracheoles which have
become spread out into a weblike form 282.
The permeability of the tracheal system shows a
gradation from the spiracle to the tracheole ending.
It is permeable to gases throughout 282; though
naturally it must be much more so where the walls
become as delicate as they are in the tracheoles. Like
the epidermal cuticle, it is probably impervious to
water, exoept in the tracheoles which are freely per-
meable. Indeed, the terminal part of the tracheoles
in many insects, both terrestrial 2?9 and aquatic 278,
normally contains a variable amount of fluid (Fig.
4, A); and it is interesting to consider why this fluid
does not extend up into the tracheae. For these
tracheal capillaries are so small (about one-third or
less of 1", in diameter) that, making certain assump-
tions, it can be calculated that the force of capillarity
in them will approximate to a pressure of ten atmo-
spheres. In other words, it will require a force of
this magnitude to keep the fluid from rising in them.
Whence can such a force be derived 1 It probably
comes from the osmotic pressure of the tissue fluids,
which is in fact of this order of magnitude (p. 37).
For if the tracheoles are exposed to salt solutions
somewhat hypertonic to the insect's blood, more
fluid is extracted from them, alld the air extends
B :'"
FIG. 4.-Tra.cheoles running to 8 muscle fibre: semi-
schematic
A, muscle at rest; terminal parts of tracheoles (shown dotted) contain
fluid; B, muscle fatigued; air extends fur Into tracheoles. (lIlodified after
Wigglesworth.)
dOWnwards towards the tissues (Fig. 4, B). And if

such solutions are replaced by hypotonic fluids, the
air retreats again higher than before. These same
changes can be brought about by the increase in
osmotic pressure which accompanies muscular con-
traction; so it is possible, that these phenomena may
on occasion have some physiological significance in
improving the supply of oxygen to those tissues whose
need is greatest 278. It is probable that osmotic
pressure is responsible also for the initial filling of
the treacheal system when the insect hatches from
the egg 290.
RESPIRATION 19
The lining of the larger tracheae is relatively hydro.
phobe (p. 6). Consequently, there is no tendency
for water to enter the spiracles. And, of course, if
the tracheoles on exposure to the air were also to
become dry and hydrophobe, capillarity would be
absent, and water from the tissues would not be
drawn along them. Perhaps that is the case in those
insects in which the tracheoles appear always to
contain air as far as they can be resolved with the
microscope.
The Diffusion Theory of Insect Respiration.
The next problem in the physiology of the tracheal
system is how oxygen is conveyed to the tissues.
Although the tracheae are permeable to oxygen
throughout their length, the abundant supply of
tracheal capillaries in and around the most active
tissues leaves no doubt that most of the oxygen enters
the organs through these endings; and the problem
resolves itself into how oxygen is conveyed from the
spiracles to the tracheoles. It was suggested long
ago by Treviranus, and later by Thomas Graham (to
whom we owe the laws of diffusion of gases), that it
must be conveyed by diffusion. But to the super.
ficial observer of these fine tracheal tubes, this ·idea
seemed so inherently improbable that various other
hypotheses were put forward, ILnd it is only within
recent years that the diffusion theory of insect reo
spiration has been generally accepted. The present
acceptance- of this theory is due to the work of
Krogh 162, wh~, taking into consideration the mean
diameter and length of the tracheae, the oxygen- con-
sumption of the insect, and the diffusion coefficient
of oxygen, calculated what difference in partial
pressure between the atmosphere and the tracheal
endings would be necessary to maintain the supply
of oxygen that was actually consumed. He obtained
the surprising result that, even in the cas~ of large
caterpillars, the partial pressure of oxygen at the
tracheal endings need not be more than 2 or 3 per
cent. below that in the atmosphere; showing that
d#fusion alone was adequate to supply the needs of
the insect.
-The same argument applies to the elimination of
carbon dioxide. The amount of carbon dioxide pro-
duced by the insect is usually rather less than the
oxygen taken in; their rates of diffusion, being pro-
portional to the square roots of their densities (V22 :
viS), are not enormously different. Therefore, since
diffusion alone will account for the supply of oxygen,
it will account equally for the elimination of carbon
dioxide. But carbon dioxide diffuses through animal
tissues, and therefore presumably through the insect
cuticle, something like thirty-five times lIlore rapidly
than oxygen 282. Consequently, since carbon dioxide
liberated in the tissues must diffuse equally in all
directions, the. amount of this gas which will escape
directly through the cuticle of the body wall (espe-
cially when this is thin) and through the walls of the
larger tracheal trunks 69, must be far greater than
the amount of oxygen that will enter by these routes.
It is probably safe to say that in most terrestrial
insects almost all the oxygen is taken in by the
tracheae; in the case of Dixippus, it has been shown
that about 25 per cent. of the carbon dioxide escapes
through the skin, and in larvae of DytisrJU8 and
Eristalis somewhat less 49.
The Function of the Spiracles. This simple
conception of tracheal respiration is complicated by
the need for retaining water; for most of the eva-
poration from an insect takes place through the
spiracles 109, 188. Burmeister discovered that each
spiracle is provided with an occlusive mechanism, the
main function of which is probably to protect the
insect from loss of water: the adult flea, which has
well-developed sphincters in the spiracles, is far more
resistant to desiccation than the larva which has
none 188. Normally, the spiracles are kept closed;
being opened only just often enough to keep the
RESPIRATION 21
insect supplied with oxygen. But when the insect
is active, or when the rate of metabolism is increased
through a rise of temperature, they are widely
dilated 115. Consequently, as the temperature rises,
the rate of water loss from an insect tends to increase
out of all proportion to the evaporating power of the'
air 109. And if the insect at rest is caused to keep
its spiracles permanently open, by exposure to
nitrogen or carbon dioxide, its rate of loss of water
is enormously increased 188.
This function of the spiracles has been termed the
'diffusion control' of insect respiration 115. It is
said to be effected chiefly by the local action of carbon
dioxide upon sensory nerve endings in the spiracles
themselves. But this cannot be the whole story, for
the spiracles are caused to open in an atmosphere of
nitrogen 115. Their opening is therefore influenced
by lack of oxygen as well as by excess of carbon
dioxide. Presumably, oxygen-want exerts its action
by the accumulation of acid metabolites, which
stimulate the respiratory centres innervating the
spiracles; and it is probable that carbon dioxide,
in virtue of its acidity, may also stimulate these same
centres.
Ventilation of the Tracheal System. This
type of respiratory system, dependent solely upon
gaseous diffusion, is adequate for most small insects,
and even large insects if they are relatively sluggish;
but it is insufficient for actively running or flying
species with a high rate of metabolism and a massive
production of energy. In these, in the larger Orthop-
tera and beetles, in bees, wasps and flies, a greater
or less degree of mechanical ventilation of the tracheal
system is superadded. We have seen that the spiral
folding of the typical tracheae renders them resistant
to collapse under pressure; but, in some cases at
least, they can be expanded and contracted in their
long axis like an accordion 282, and their capacity
reduced by as much as 20 to .30 per cent. 69. In other
cases they are not round in cross-section but elliptical
(Dytiscus larva) or even ribbon-like (in the thorax of
Muscid flies) and will then readily collapse when the
'pressure around them is increased. Or they may
have thin-walled cavities at intervals along their
course (as in Melolontha), or large dilatations (' air
sacs ') which occupy a great 'part of the body cavity
(as in many Orthoptera, Hymenoptera, &c.) (Fig.
5, B) 39.
There is little doubt that, as Treviranus suggested,
the main function of these collapsible traoheae or air
saos is the ventilation of the respiratory system. Like
the ventilation of the lungs of vertebrates, the filling
and emptying of these air saos is seoondary to the
respiratory movements of the rigid body wall whioh
enoloses them. When the body wall is in the in-
spiratory position they are widely open; in the ex-
piratory position many of them are oollapsed and
empty 89. In addition they are doubtless ventilated
to some extent by general movements of the body
-for example, during flight; and where they lie in
rigid portions of the body, suoh as the head, they are
ventilated, as Graber pointed out, by the transmitted
pressure of the blood.
The result of this process is that the greater part
of the traoheal syst~m is kept filled with a gas whioh
approximates in oomposition to the outside air; but
the aotual supply of oxygen to the tissues again takes
plaoe by diffusion along the tracheal branohes given
off by the air saos. This, of oourse, is analogous to
what happens in vertebrates, where only the upper
parts of the respiratory tract are ventilated me-
ohanically, exohanges in the alveoli of the lung being
dependent on diffusion. The extent to which the
traoheal system is ventilated mechanioally varies
enormously in different inseots, and in the same insect
acoording to its physiologioal state; but in many
oases the efficienoy of the prooess compares very
favourably with the ventilation of the human lungs.
RESPIRATION 23
For instance, in Melolontha the respiratory system
may be emptied during expiration of about one-third
of its total capacity 69 (the total capacity being 39
per cent. of the body volume 1), and in the larva of
Dytiscus of about two-thirds 162. (In quiet respira-
tion in man, the lungs are emptied of about one-
seventh of their total capacity; the most extreme
degree of ventilation empties about two-thirds.)
(In addition to their ventilating function, the
tracheal air sacs of insects sometimes serve a purpose
like the air-containing sinuses in the facial skeleton
of mammals, or the pulmonary air sacs in the bones
of birds; thus, by extending into the massive head
and mandibles of certain beetles, they permit an in-
crease in the skeletal bulk without adding to the
insect's weight 282.)
The respiratory movements of insects are brought
about by a special musculature which varies greatly
from one group to another 9. As a general rule the
movements are confined to the abdomen, but in
Dytiscus and Hydrophilus ventilation is maintained
by aspirating movements of the metathorax, the
abdominal movement apparently being passive 40.
In most cases expiration is active and inspiration
passive, though in the larva of Aeschna and in the
grasshopper expiratory muscle~ also are present 282.
The movements may consist of dorso-ventral :Batten-
ing movements (Orthoptera, Coleoptera) or longi-
tudinal telescoping movements (Diptera, Hymen-
optera).
The simplest interpretation of the!le movements is
that they are ventilating all parts of the tracheal
system alike. But if, during the respiratory con-
traction, the spiracular valves in different parts of the
body open and close independently of one another, it
is possible that a directed stream of air will be driven
through the tracheal system-certain spiracles serving
always for inspiration, others for expiration-an
arrangement which would serve to enrich the air in
3
the inspiratory section of-the system. The idea that
such a circulation of air does occur has often been
put forward 282, but only in recent years has it
been convincingly demonstrated. Thus, it has been
shown that if the four anterior pairs of spiracles in
the locust (Schistocerca) are enclosed in one gas
chamber, and the six posterior pairs in another, air
is actively transported during respiration from the
anterior to the posterior chamber 90, 181. The under-
lying principle of this mechanism is that the rhyth-
mical nervous impulses which control the ventilating
movements are co-ordinated with impulses which
bring about the synchronous movement of the
spiracles-some of these opening during inspiration,
others during expiration 90: But before looking
further into this point, we must consider the control
of the ventilating movements themselves.
The Regulation of Respiration. The respira-
tory movements are effected by impulses from nerve
centres. These centres are of two kinds 239: so-
called primary respiratory centres which lie in the
segmental ganglia and control the movements of their
own segments (for the isolated segments of the
abdomen may perform respiratory movements) 9, and
secondary centres which have an overriding action
and control the movements in the whole insect. One
might expect the secondary centre to lie in the head,
but this seems never to be the case: decapitation
produces only a temporary and uncertain effect on
respiration 282. In the few insects that have been
studied, it is situated in the prothorax 239.
The isolated g_anglia of insects which possess a
mechanical ventilation of the tracheal system (Dytis-
CUB, Aeschna) show rhythmical changes in electrical
potential of the same frequency as the respiratory
movements of the intact insect 2. There must,
therefore, be a spontaneous respiratory rhythm in-
herent in the nerve centres. But the activity of the
centres, both in rate of rhythm and depth of move-
RESPIRATION 25
ment evoked" may be influenced by external stimuli,
either nervous or chemical 90; and during times of
rest the ventilating respiratory movements may
cease altogether 115. There is evidence that during
the onset of activity-flight, for example-the res-
piratory movements may sometimes be initiated. by
nervous stimuli alone 90 (for any stimulus will cause
an increase in respiration in insects) 239; but they
can also be initiated by chemical stimuli alone 116 ;
and, although there must be endless variation in
different species, perhaps these play the greater part
in the normal life of the insect.
Either a lack of oxygen 9 or an excess of carbon
dioxide 239 may stimulate the respiratory centres and
cause hypernoea; in some insects an excess of oxygen
may cause a prolonged apnoea 9. Now excess of
oxygen will not diminish the production of carbon
dioxide; nor will a lack of oxygen increase it. If,
therefore, carbon dioxide were the 80le respiratory
stimulant, respiration should not be influenced by the
tension of oxygen. But there is no doubt that carbon
dioxide can control respiration: in DixippU8, the
activity of the secondary or prothoracic centre is
stimulated by carbon dioxide tensions of 0·2 to 3 per
cent.; the primary centres respond to 12-15 per
cent. 239; and in the Periplaneta and other insects,
ventilation begins, even in the insect at rest, at a
carbon dioxide tension of 10 per cent. 115 Since it is
reasonable to assume that the chemical factor at
work is the same in all these cases, it seems probable
that this factor is the acidity in the nerve centres-
due either to an excess of carbon dioxide, or to acid
metabolites accumulating in the absence of oxygen 239.
But some authors believe that there may be receptors
(which have not been demonstrated histologically)
in the tracheae, which respond to the gas these
contain 239.
We have seen (p. 20) that in thin-skinned insects
much of the carbon dioxide diffuses through the
26 INSECT PHYSIOLOGY"
general surface of the body, and therefore cannot
serve as a respiratory stimulus at all. Thus we find
that mosquito larvae, and the larvae of Aesckna and
Dytiscus, are driven to seek air at the water surface
by oxygen want and not by carbon dioxide ex-
cess 162, 282. On the other hand, the leg movements
in the aquatic bug Oorixa, which direct a stream of
water over its air store (p. 28), are called forth by
the accumulation of carbon dioxide; while the same
insect is caused to rise to the surface when the air
store is reduced to a given size 32.
We are now in a position to revert to the question
of the stream of air directed through the tracheal
system by the co-operation of ventilatory and
spiracular movements. In the light of recent work
there is no reason to doubt that such a directed
flow does often occur. But its existence implies that
the spiracular opening is being centrally controlled
by an extremely complex nervous mechanism which
synchr~mizes this movement with the ventilation 90.
Now we have seen that the spiracles are normally
chemically controlled 115. This chemical control may
be quite local in its action; in the sense that those
spiracles which serve the most active parts of the
body may show rhythmical opening and closing while
the remainder of the spiracles stay closed 180. Con-
sequently, it must always be a matter of difficulty to
decide whether, in a given insect, the occurrence of
a directed stream of air through the tracheal system
is a purposive movement co-ordinated by a higher
respiratory centre, or whether it is a purely accidental
result of the simultaneous occurrence of a ventilat-
ing rhythm determined by the state of the higher
FIG. 5.-Some types of respiratory system in insects:
schematic
A, simple alUU!tomoslng tracheae, with sphincters in the spiracles (p. 16);
B, mechanically ventilated air sacs developed (p. 22); C, metapneustic
respiration: only terminal spiracles functional (p. 28); D, tracheal system
entirely closed: cutaneous respiration (p. 29); E, the same with abdominal
tracheal gills (p. 30); F, the same with rectal tracheal gills (p. 30).
FIG. 5.
28 lNSECT PHYSIOLOGY
respiratory centre, and a spiracular rhythm deter-
mined by the local needs of different parts of the
body 180, 181. To decide between these alternatives
will need very careful study in each insect.
Respiration of Aquatic Insects. The majority
of insects that are aquatic in the adult stage breathe
gaseous air like the terrestrial forms; but they
develop special structural modifications to enable them
to do so. Since all the spiracles cannot be exposed
to the exterior, there is a tendency for the respiratory
function to become concentrated at one end of the
body. In Hydrophilus, the mesothoracic spiracles
are the most important; in DytiSCU8 and its allies
and in the aquatic Hemiptera, the last abdominal
spiracles predominate 84. In many larvae (DytiSCU8,
Culicidae and many other Diptera) the other spiracles
have become obliterated, or at least functionless, and
the insects breathe entirely through the terminal
spiracles (' metapneustic' respiration) (Fig. 5, 0).
In all these cases, in the antennae of Hydropkilu8, in
the abdominal fringes of hair in Notonecta, in the
elytral surface itself in Dytiscus, and around the
tracheal orifices of the metapneustic larvae, there are
hydrofuge structures that enable the air-containing
regions of the body to break through the surface film
and obtain communication with the atmosphere 282
(see p. 6).
In many cases (mosquito .larvae~ Dytiscus larvl;Le,
&c.) the tracheal trunks are so capacious that they
serve as stores of air, and enable the insect to remain
submerged for a considerable time. In other insects,
a store of air is carried beneath. the elyt_ra (in Dytiscus
&c.) or over the ventral surface of the body (in
Notoneda, Oorixa, &c.) by means of the hydrofuge
surfaces and hairs already discussed (p. 6). These
external air stores have several functions: (1) They
have an important hydrostatic function in enabling
the insect to reach the water surface in the correct
position to take in oxygen 282. (2) They provide a
RESPIRATION 29
store of oxygen which is gradually used up by the
submerged insect; for example, the oxygen in the
elytral air of Dyti8CU8 may fall from 19·5 per cent.
at the moment of diving, to 1 per cent. or less in
three or four minutes 282. (3) They provide a mech-
anism for obtaining dissolved oxygen from the water,
and so function as a kind of physical gill 76. For the
invasion coefficient of oxygen between water and air
is more than three times as great as that of nitrogen.
Consequently, as the partial pressure of oxygen in
the air store becomes reduced, equilibrium will be
restored by the diffusion of oxygen inwards, rather
than by the diffusion of nitrogen outwards. Of
course, some nitrogen will diffuse out, but as long as
any remains undissolved the process can go on, and
the insect can extract dissolved oxygen from the
water. This mechanism is of more or less value to
all aquatic insects that carry air stores; and small
forms like Corixa can obtain enough oxygen in this
way even at summer temperature so long as they do
not swim actively. The stI·iking effect of the process
may be shown by quoting a single experiment: Noto-
necta lived five minutes submerged in water saturated
with nitrogen, thirty-five minutes in water saturated
with oxygen, but six hours in water saturated with
atmospheric air-:-the air store, in each instance, being
first charged with the dissolved gas 76.
By the provision of air stores which cannot be 're-
placed by water, a few insects have become indepen-
dent of the atmospheric air; but this has happened
very rarely 282. On the other hand, by the obliter-
ation of all the spiracles of the respiratory system,
and the development of gills, many insect larvae have
become completely adapted for the respiration of
dissolved oxygen 64. In many cases, the general
surface of the larVa, richly supplied with fine tracheae,
provides the respiratory surface (in Chironomu8,
Simulium, Corixa, &c.) (Fig. 5, D). Sometimes there
are specialized plate-like projections rich in tracheae
(Ephemeridae, Plecoptera) (Fig. 5, E); or papillae
from the anus, consisting of little more than massed
tracheal branches covered by a thin cuticle (Elmis,
Col.); or' a network of tracheae inside the rectum
(as in the larvae of Anisopterid dragon-flies) (Fig. 5,
F); or rigid cutIcular filaments containing air, which
do not collapse on drying, and can therefore serve for
respiration both in water and on land (as in the pupa
of Simulium, and other Diptera from mountain
torrents 215).
In some cases the oxygenation of these organs is
ensured by a highly developed respiratory centre:
the gill plates of Ephemeridae are violently agitated
in water poor in oxygen 9, and the rectum of Aeschna
is ventilated by abdominal movements controlled by
primary and secondary respiratory centres (p. 25) 239.
It has been shown that the partial pressure of oxygen
in the closed system of these insects is aways less
than its tension in the surrounding medium, and that
of carbon dioxide greater; it is therefore generally
believed that the exchange of gases through these gills
is effected solely by diffusion 282.
In addition to these undoubted gills, there are, in
many aquatic insect larvae, tubular out-growths from
the body surface which may contain blood only, or
blood with a rather sparse supply of tracheae. These
st~uctures are called blood gills; but what experi-
mental evidence exists seems to point against their
having any but a subsidiary function in respira-
tion 114, 253; though in the present state of know-
ledge it would be unwise to generalize 266. In one
case, that of the anal papillae of the mosquito larva,
they have been shown to be chiefly water-absorbing
organs (p. 68) 285.
We have seen (p. 28) that the external air stores
carried by aquatic insects help to control their equili-
brium in the water. The air in the tracheal system,
also, must often be of some importance for this pur-
pose; a.nd in the pelagic larva of Oorethra there are
RESPIRATION 31
developed four large air sacs which function solely
as hydrostatic organs. This larva can adjust its
specific gravity to that of.the water it is in by varying
the capacity of these air sacs-apparently by some
active change which causes expansion or contractio~
of their cuticular walls 282.
Respiration of Parasites. The internal para-
sites of insects, which live a semi-aquatic life within
the blood and tissues of their host, naturally show
many of the respiratory adaptations possessed by
aquatic larvae. In the young stages of zp.any of them
the tracheae do not contain air, and oxygen simply
diffuses from the blood of the host into the blood of
the parasite-a~ it does in the young aquatic larva
of Ckironomu8 (Dipt.) and Acentropua (Lep.), &c.
Later, when air has appeared in the tracheae, these
commonly supply a rich network of branches to the
skin-as they do in the older aquatic larvae of
Simulium, Ckironomu8, &c. There are often out-
growths from the body surface, usually from the tail,
which recall the gill-like organs of aquatic forms.
Sometimes these structures are well supplied with
trache!J.e (as in the larva of Cryptockaetum (Agromy-
zidae: Dipt.» 251 or with circulating blood (as in
Apantele8 (Braconidae: Hym.» and are of proved
importance in respiration; but in most cases their
respiratory significance is very doubtful and their
function is problematical 252. Finally, the meta-
pneustic type of respiration common in aquatic
insects is seen in most Dipterous parasites (Tachi-
nidae, &c.), which pierce the body-wall or the large
tracheal trunks and breathe the atmospheric air. The
habit of attaching themselves to the tracheal tubes.
recalls the fact that certain aquatic larvae (of the
beetle Donacia and the mosquito Man8onia) possess
specially modified respiratory siphons which they
insert into the air-containing tissues of Iltquatic plants,
and are thus able to remain permanently beneath
the water surface 282.
CHAPTER III
THE CmCULATORY SYSTEM AND THE BLOOD
N the insect body there is only one tissue fluid,

Icavity,
the blood or haemolymph, occupying a single
the haemocoel, and freely bathing all the
tissues. With few exceptions, there is only one
blood-vessel, which runs along the mid-line of the
back. The posterior segment of this pulsating
vessel, the 'heart' , is provided with a series of
valved openings or ostia through which the blood can
enter; the anterior segment, the' aorta', is a uniform
contractile tube. After passing through the brain
above the oesophagus, the aorta ends more or less
abruptly Ill; and from this point the blood simply
percolates slowly backwards through the tissues.
But in many insects it is still subject to some direc-
tion: the aorta may discharge into definite vessels
carrying the blood in different directions 40; the
a:Q.tennae and limbs are often divided by longitudinal
membranes, the blood entering the limb on one side
and leaving on the other; and the abdominal cavity
in some insects is divided by two horizontal mem-
branes, the dorsal and ventral diaphragms, into three
sinuses: the pericardial sinus containing the heart,
the perineural sinus containing the nerve cord, and
the visceral sinus between. The perineural sinus
tends to conduct the blood to the posterior end of
the abdomen before allowing it to return to the
heart; hut it is perforated and incomplete laterally,
so that some of the blood circulates transversely
across the abdomen below it. The dorsal diaphragm,
32
THE cmCULATORY SYSTEM 33
when present, is also fenestrated, and thus allows
the blood to enter the pericardial sinus throughout
its length (Fig. 6).
The Mechanism of the Circulation. In insects
with a rigid cuticle, the blood in the 'body cavity,
when the insect is at rest, is at atmospheric pressure
(Odonata 232) or even less (DytiscU8 40, Apia 91). In
the more soft-bodied insects, the general pressure
may be raised (to 40 or 50 mm. of water in the larva
of Aeschna, or even more during exertion 232) by the
muscular tension· of the body-wall. The pressure
may be increased, also, during the ventilation of the
tracheal system; and during those special acts which
are brought about by the displacement of the blood,
such as the expansion of the wings in emerging adults
(p. 13), the general pressure may be maintained for
a prolonged period at a high level (75 mm. of water
in the dragonfly 232).
But, like the blood in the vena cava of mammals,
the blood of insects is always aspirated, into the heart,
during diastole, under a negative pressure: a pressure
less than that existing in the general body cavity.
This force is due partly to the elastic muscular walls
of the heart itself73, partly to the elastic traction of
the dorsal diaphragm which is tied to the lower wall
of the heart 40, and partly to the contraction of the
muscle fibres' (the aliform muscles) which commonly
occur in the substance of this diaphragm 73. During
systole, a weak positivI') pressure develops and the
blood is driven forwards.
In some insects, such as the larvae of Nematocera 73
and the Ephemeridae 190, the valves of the ostia have
become so modified as to form interventricular valves;
these divide the heart into a succession of chambers
through which the blood can How only forwards. But
this arrangement is the exception; in most insects
the heart is patent throughout its length and acts
as a whole 101, the blood being carried along it by
peristaltic waves. As a rule, such waves start at the
hind end of the hea:cti and pass forwards; but in
many insects 101, 211 the direction of beat may be
periodically reversed, and the blood then escapes
into the abdominal cavity through the relaxed ostia.
Sometimes beats may arise at a number of points
B st'
FIG. 6.-A, insect with fully developed circulatory system :
schematic; B, transverse section of thorax of the same;
C, transverse section of abdomen. Arrows indicate
course of circulation. (Based largely on Brocher.)
a, aorta; apo, accessory pulsatile organ of antenna; d, dorsal diaphragm
with aliform muscles; h, heart; n, nerve cord; 0, ostia; pc, perlcardlal
sinus; pn, perineural sinus; po, meso- and metathoracic pulsatile organs ;
B, septa dividing appendagea; fl, ventral diaphragm; "B, visceral sinus.
simultaneously and extinguish one another when

they meet 73.
Thus the circulation of the blood is secured prim-
arily by the work of the heart, which aspirates it
from the abdominal cavity and pumps it forwards
to the head. In the thorax, accessory pumps are
THE CIRCULATORY SYSTEM 35
often present 40. These aspirate blood from the
thoracic cavity, through certain of the wing veins,
and return it through connecting vessels either to the
aorta itself (Dytiscus, Aeschna larva, Sphinx) 39 (Fig.
6, A) or to the body cavity again (Tabanus, Dipt ..;
Vespa 39, Apis, Hym. 91). Accessory pumping organs
may irrigate the antennae (Periplaneta, Hymenop-
tera, &C. 91 ) and the legs (Hemiptera) 111. But in
many insects these structures are wanting, and the
circulation through the appendages is thep effected
(i) partly by their own muscular movements, (ii}
partly by the pressure,changes in the abdomen brought
about by the respiratory movements (Aeschna larva 40,
Ephemeridae larvae IDO)-in which case the pulse
beats in the leg synchronize not with the heart-beat
but with the respiratory contractions, and (iii) partly
owing to the fact that the efferent stream (in the
posterior chamber of the limb) communicates with
the perineural sinus, where the pressure is higher,
while the afferent stream (in the anterior chamber)
communicates with the perivisceral sinus, where the
pressure is lower: the blood in the limb is thus
subject to the aspirating action both of the. heart
and of the pulsatile organs of the thorax 40 (Fig. 6,
B). Finally, in those insects in which the ventral
diaphragm is well developed (many Orthoptera,
Hymenoptera, Lepidoptera), it may contain muscle
fibres, and by means of wave-like contractions may
actively assist in driving the blood backwards and
laterally 40.
Control of the Heart-beat. If the heart of the
cockroach may be taken as a normal example, the
heart of insects has a very complicated nerve supply 3.
Two lateral nerves, containing ganglion cells, and
apparently cQnstituting an intrinsic cardiac ganglion,
run along its entire length, and are connected in front
with the ganglia of the visceral nervous system. In
addition, the heart receives sensory and motor nerve
fibres from the segmental ganglia of the ventral chain.
There is no doubt, therefore, that the heart enjoys
a large degree of nervous control; experimentally,
faradic stimulation in the region of the neck gives
rise to accelerating impulses which reach the heart
both by the lateral nerves and by the segmental
branches from the ventral cord 241. What is less
certain is whether the automatism or spontaneous
pulsation of the heart is a property of the muscles
(myogenic) or of the nerves (neurogenic). An im-
portant factor in the initiation of each beat appears
to be the tension in the muscular wall of the heart,
due either to the contraction of the aliform muscles
(which, in a given segment, may precede the con-
traction of the heart itself) or to the constant elastic
tension which these structures exert 73. lfthe aliform
muscles are cut through, the heart no longer beats 73.
But it does not necessarily follow from this that the
heart-beat is myogenic. It has been found that the
change in rate of the heart-beat at different temper-
atures shows a 'temperature characteristic' or
, critical thermal increment' which agrees remarkably
well with that of other vital activities which are con-
trolled.by the rate of discharge from nerve centres 241 ;
and for those who regard such evidence as carrying
weight, this suggests a neurogenic origin. Whatever
the nature of the automatism,it is remarkably sensitive
to carbon dioxide: the hearl-beat in aquatic larvae
is at once arrested in water saturated with this gas 88.
Composition and Properties of the Blood.
The blood of insects may be a clear colourless fluid;
but more often it is tinged with green or yellow
(p. 81). The specific gravity varies from 1·03 in
the larva of Deilephila (Lep.) 119 to 1·045 in the larva
of ApiB 22. The reaction is usually very slightly acid:
pH = 6·4 in Deilephila larva 119, 6·6-6·8 in Bomhyx
mori 68, 6·83 in ApiB larva 22; but it may differ
slightly in the two sexes 68 and may vary with the
age of the insect. Thus, in Bomhyx it becomes more
alkaline at each moult 68; and in Apis 22, Deilephila 119
THE BLOOD 37
and other insects 179 it is more acid during the pupal
stage. The blood is moderately well buffered; but
the bicarbonate is very low 74 and the buffering
appears to be due to the phosphate and, presumably,
the protein. The molecular concentration, expressed
as t:,. ranges from 0·56 in Saturnia larva 74 to 0·78-0·81
in Deilephila larva 119, and 0·86-0·87 in Apislarva 22;
but the osmotic pressure can, apparently, become
enormously increased temporarily by the accumula-
tion of unoxidized metabolites during oxygen want 1I78.
On the other hand, those few insects which have.
colonized sea water appear to be homoiosmotic: the
molecular concentration of their blood is unaffected
by the outer medium 285. Of the total molecular
concentration, a surprisingly small part, sometimes
not more than 4·2 per cent. and rarely more than
15 per cent., is chloride 74; whereas the phosphate
content, as compared with vertebrate blood, is very
high, as are, also, the magnesium and calcium 38,
But the most striking feature is the high content of
amino acids 22,74, which may be twenty or thirty
times as great as in human blood, and form nearly
15 per cent. of the total nitrogen 59. In addition,
there is a considerable amount of residual nitrogen
in peptide form 119, while proteins (albumen and
globulin) are present in much the same amounts as
in human plasma. The glucose level is of the same
order as in mammals 22, 119. Urea is always present
in small quantities 67. Fat, which occurs largely as
a very fine emulsion, varies with the state of diges-
tion and nutrition 110.
The blood of many insects (Apislarva 22, &c.) does
not clot; wounds are simply-closed by a plug of cells.
In others (cockroach, &c.) it clots very readily. The
blood-cells send out fibrillar pseudopodia which inter-
lace with one another to form a living network within
which the fibrin filaments separate out; but the cells
are said not to be necessary for clotting to take place,
nor is calcium needed for the process 200.
Cytology of the Blood. The blood. cells or
haematocytes of insects, as seen in preparations,
present an extraordinary di~"ersity of appearance;
but this seems to be due to the protean forms which
they can assume rather than to the abundance of
distinct types. They multiply and grow in the body
cavity throughout the life of the insect, appearing
first as small darkly staining forms termed 'pro-
leucocytes' 132, 211, which are often seen dividing and
are not yet capable of phagocytosis. As they grow
they become pyriform or spindle-shaped and will
ingest dead bacteria, Indian ink, tissue debris, &c, ;
they are then called' phagocytes '. Their appearance
naturally varies very much with their content, and
sometimes separate names have been given to these
phases 137; moreover, they have a habit of spreading
themselves out in stellate form upon flat surfaces, such
as the basement membranes, their aspect becoming
totally changed 286.
These haematocytes are by no means all circulating ;
indeed, the majority adhere to the surface of the
tissues; they collect particularly along the sides of
the dorsal vessel, often in definite clumps termed
, phagocytic organs' 62, The larvae of Chironomidae
present a complete series in this respect: in certain
genera only free haematocytes are present; in others,
both haematocytes and fixed phagocytic cells occur;
and in yet others there are no circulating (lells at all
but only phagocytic tissue 171. The haematocytes
becOIpe far more 'numerous during moulting 286 and
metamorphosis 208, when they may play a part in
removing the dead cells and tissues: during the meta-
morphosis of Muscids, these phagocytic cells (the
'K6rnchenkugeln' of Weismann), stuffed with the
products of histolysis, are exceedingly conspicuous.
The phagocytic haematocytes will ingest, and in some
cases destroy, living bacteria introduced into the
blood 137, They collect at the site of wounds and form
a closing plug which forms the basis for the subsequent
THE BLOOD 39
healing process. They will congregate in great
numbers around foreign bodies 172, 218 and certain
parasites, walling these off in a discrete capsule;
sometimes the cellular nature of this capsule per-
sists 135, but more often the cell bodies become con-
verted into homogeneous membranes from which the
nuclei disappear; and it is possibly in this way that the
connective tissue membranes which cover the organs
of insects are formed 172. In some insects, haema-
tocytes laden with conspicuous granules (granular
leucocytes) 132 are present in the blood; although
these are no longer phagocytic they seem to represent
a stage in the life history of these same cells.
Besides this type of cell with all its varied powers,
there is, in most insects, another type which is equally
distinct. It has a rounded or oval form, the cyto-
plasm is generally eosinophil, and it takes no part
in phagocytosis 132 nor in the formation of encapsu-
lating membranes 172. In appearance these cells
are like diminutive oenocytes; and on this account
they are often called 'oenocytoids'; but they do
not seem to bear any relation to the true oenocytes 286.
Their function is quite unknown: they increase in
number during moulting 286, like the phagocytes; and
they collect around foreign bodies in a zone peripheral
to the phagocytes themselves 172.
Finally, in certain Hemiptera, there are blood-cells
laden with fat globules, , adipoleucocytes " and others
charged with wax; but the relation of these to the
types described above is uncertain 132.
The Functions of the Blood. (i) Mechanical.
The blood plays an important part in transmitting
pressure from one region of the body to another: the
hatching of many insects from the egg (p. 13), the
rupture of the old skin at moulting (p. 12), the ex-
pansion of the wings in the newly hatched adult insect
(p. 13) and many other movements are, brought about
by the localized pressure of the blood~
(ii) In Nutrition. The blood conveys the nutrient
.d.
40 INSEct PHYSIOLOGY
materials to the tissues, and the waste products to
the excretory organs. The precise part that it plays-
in these functions is not really understood, but it is
probable that the composition of the blood may vary
enormously with the state of nutrition and, notably,
during moulting and metamorphosis when many of
the larval tissues are breaking down and the cells of
the fat body are yielding up their protein and fatty
contents. The blood must also be regarded as an
important reserve of food material: in the larva of
Deilephila, during fasting, the protein in the blood
is rapidly consumed; apd during pupal life in the
same insect more 'than half the total energy meta-
bolism is effected at the expense of the blood 119.
(iii) In Immunity. The phagocytic blood-cells
provide perhaps the chief mechanism for protecting
the insect from bacterial invasion 64, 137 ; but humoral
immunity, both natural and acquired, does occur 1i4.
The discussion of these matters would take us too
far afield.
(iv) In Respiration. We have already seen that
oxygen is ordinarily conveyed directly to the tissues
by the tracheal system. But many cells are separated
from the nearest tracheal tubes by an appreciable "
space; sometimes, notably in the pupa, organs may
be entirely devoid of tracheal supply; and in some
aquatic insects the tracheal system is completely
filled with fluid. Moreover, the tracheal system,
throughout its length, is permeable to oxygen, which
must therefore escape into the blood. Under all
these circumstances 282, the blood acts as a carrier of
oxygen. There are, also, here and there among
insects, special arrangements of the tracheal system
which seem to be designed to aerate the blood: a
convoluted tract in the aorta of the honey-bee is
richly supplied with tracheae 01; the posterior region
of the heart in the metapneustic larvae of Nematocera
is often surrounded by a basket-work oftracheoles 282;
and in the thorax of N epa is a peculiar organ so
THE BLOOD 41
richly supplied with tracheae and bathed with blood
as to suggest that it may be a kind of tracheal lung 111.
In view of these relations the question naturally
arises whether the blood of insects contains chemical
carriers of oxygen comparable with the haemoglobins
and haemocyanins of other animals. With a single
exception, this has not yet been proved to be the
case. In those insects, such as the larva of the honey-
bee, in which the properties of the blood have been
most carefully studied, it has not been found to take
up more oxygen than can be accounted for by physical
solution 22; but it must be admitted that such in-
vestigations have not yet been very extensive. The
single exception is in the larvae of certain Chirono-
midae, which contain haemoglobin in the blood in
free solution-that is, not in corpuscles. The affinity
of this haemoglobin for oxygen is, however, very
different from that of mammals. At all ordinary
tensions of oxygen it remains fully saturated; and
it only begins to liberate its oxygen when the pressure
of this gas is reduced to I per cent. of an atmosphere
or less; the precise level varying, in different
species 1U.. In other words, it is capable of acting
as a carrier only under conditions of extreme paucity
of oxygen and seems, in fact, to be a special adaptation
to life in poorly oxygenated waters-the larvae that
contain haemoglobin being much less susceptible to
oxygen want"282.
We have seen, also, that the blood probably plays
a considerable part in the carriage of carbon dioxide
from the tissues (p. 20). But here again there seems
to be no chemical provision for its transport; the
bicarbonate and the carbon dioxide capacity of insect
blood are both very low 22, 74.
CHAPTER IV
DIGESTION
NSECTS feed upon almost every type of organic
Iothers
substance found in nature some on plants,
259:
on animals; some on ·the sap of plants or the
tissue fluids of animals, others on foliage, or flesh,
dry timber, or hair and feathers; some on fungi or
the live and dead bacteria in the excrement of animals,
others on the sterile juices of living forms. Their
feeding mechanisms, and the structure and chemistry
of their digestive system present, therefore, the most
extraordinary variety, and it becomes singularly
difficult to sift out the general principles in the phy~io
logy of their nutrition.
The Alimentary Canal. With the exception of
the earliest stages of some parasitic insects, which
absorb nutriment through the general body surface £51,
all insects take their food into an alimentary canal.
This consists always of three parts: the fore-gut,
mid-gut and hind-gut. The fore-gut and hind-gut are
both lined with cuticle; in the mid-gut the cells are
freelyexposed; from which it follows that secretion of
digestive juices can occur only in the mid-gut, and
here, also, at least the greater part of absorption un-
doubtedly takes place. In many insects the secretion
of the mid-gut is supplemented by that from the
salivary glands, mixed with the food before it is
swallowed.
Fig. 7 shows some of the commoner types of intes-
tinal svstem, and the different uses to which their
parts are put. In the most primitive insects, and in
42
:. G
FIG. 7.-Types of alimentary system in insects: schematic
T.
Fore·gu_t and hlnd·gut indicated by a heavy ine internally. Explanation
in text. (0 modlllec1 after Weber.)
many larval forms (Diptera.-Nematocera, Lepidop-
tera, Tenthredinidae and many Coleoptera) (Fig. 7,
A), the fore-gut or oesophagus has no other function
than to conduct the food into the mid-gut; from
which it is passed on, often more or less continuously,
to the hind-gut. But in very many insects (Dermap-
tera, Orthoptera, Isoptera, Odonata, Hymenoptera,
many Coleoptera), the hinder part of the fore-gut is
dilated to form a capacious crop (Fig. 7, B). Here
the food is stored before being transmitted, in small
quantities at a time, to the mid-gut. It is, however,
not only stored but digested, being acted upon always
by the salivary secretion, and, at least in the Orthop-
tera, by the digestive juices passed forward from the
mid-gut U7. In the higher Diptera a crop for storing
food is again present (Fig. 7, C), but it takes the form
of a diverticulum connected to the fore-gut by a
narrow duct. The food in this crop is mixed only
with saliva, and it suffers very little digestion until
it is transferred t9 the long coiled mid-gut. In the
Lepidoptera, and in many Diptera (Culicidae, Taban-
idae 281) (Fig. 7, D), the ingested food goes straight
to the mid-gut and is there both stored and digested;
the 'crop' has all but lost its function as a food
reservoir, and is used chiefly (in many Lepidoptera,
solely) to receive the air swallowed by the insect
when it emerges from the pupa (p. 13). In the fleas
(Siphonaptera) and sucking lice (Siphunculata) the
crop has been lost entirely (Fig. 7, E), and the food
is taken straight into a voluminous stomach where it
remains until digestion is complete.
In many Diptera the mid-gut consists of several
segments characterized by differences in the epithe-
lium; the more anterior being concerned probably
only with absorption, the more posterior with diges-
tion 277. The value of this arrangement when the
insect takes in much fluid with its food is obvious;
for much of the water .and many of the assimilable
constituents are removed before the food reaches the
DIGESTION 45
digestive juices, and consequently these juices do not
suffer unnecessary dilution. In many Heteroptera,
this arrangement has gone a stage further; there is
once again a capacious crop in which the meal is
received, but it is a crop composed of mid-gut. In
the blood-sucking forms (Oimex and Rhodniu8) (Fig.
7, F), the food is not at all digested in this part of
the mid-gut, but is merely concentrated by the re-
moval of fluid, and the concentrated product passed
on for digestion to the long narrow intestine 288. In
these insects the fluid that is absorbed passes into
the blood and is then excreted by the Malpighian
tubes 176, 283; but an advance on this mechanism has
been developed by many Homoptera (Coccidae, &c.).
These feed on the juices of plants, and therefore receive
a great excess both of water and of sugars. But instead
of the superfluous fluid being taken into the blood
and then eliminated by the Malpighian tubes in the
manner just described, a dilated loop of the fore-gut,
the 'filter chamber', with very delicate walls, is
invaginated into the rectum .(Fig. 7, G); the un-
wanted fluids are here absorbed, or perhaps simply
filtered, directly into the terminal part of the hind-
gut; and so come to be disc4arged again to the
exterior as honey dew, manna, and such-like pro-
ducts 268. Many modifications of this arrangement
have been described 268.
Another special modification of the gut is met with
in the larvae of Hymenoptera-Apocrita, and in the
'larvae of Myrmeleon and other Neuroptera (Fig. ,7,
H). In these the mid-gut remains unconnected with
the hind-gut until the time of pupation. Conse-
quently, the stomach becomes enormously distended
with the undigested residue of the food, which is not
evacuated until just before pupation 19.
The Proventriculus. In nearly all insects there
are special structures where the fore-gut and the m~d
gut join. There is nearly always a muscular sphincter
by which t1;te contents of the two regions can be kept
separate; and there is often a discrete organ known
as the proventriculus. The proventriculus is a different
morphological entity in different groups, sometimes
being composed solely of fore-gut, while sometimes
both fore-gut and mid-gut contribute to it. Its
function also varies. In the Blattidae it is a powerful
muscular organ or gizzard provided with rows of
massive teeth, which seize and triturate the solid par-
ticles of food in the crop 79. In many Hymenoptera
it is a similar type of organ though with smaller teeth,
but it has not been proved to have a grinding
function 237. In the flea it is lined with long needle-
like teeth directed backwards; during digestion it
contracts rhythmically, driving the teeth backwards
into the mid-gut; at the same time the mid-gut
contents are driven up against it by anti-peristaltic
waves, and in this way the blood corpuscles are
pounded and disintegrated 1188.
In most insects the fore-gut is invaginated to some
extent into the mid-gut, and this invagination, sur-
rounded by the anterior or cardiac region of the mid-
gut, is also commonly referred to as the 'proventri-
culus '. The invaginated fore-gut is called the
'oesophageal valve', though it seems never to
function as a valve (if, by that word, is intended a
mechanism which automatically allows movement in
one direction only), for the occlusion of the passage
between-fore-gut and mid-gut is secured always by a
sphincter muscle 280, and the invagination of the fore-
gut has quite another significance.
The Peritrophic Membrane. The gut of in-
sects contains no mucous glands; the boluses and
hard particles of food are therefore not lubricated as
they are in the intestine of vertebrates; and the
epithelial cells obtain protection in another way. As
was first observed by Lyonnet, they are separated
from the gut contents by a delicate membrane, com-
posed of chitin mixed perhaps with some protein 277,
known as the peritrophic membrane. This membrane
DIGESTION 47
is freely permeable to digestive enzymes and to the
products of digestion; experimentally it has been
shown to be permeable to all dyes save those with
the largest colloidal particles 65. In its primitive state
the peritrophic membrane is a somewhat_ indefinite
affair formed by the condensation over the surface
of the food of certain constituents in the mid-gut
secretion; and it arises in this way to a greater or
less extent in many existing insects, where it is made
up of a number of concentric layers, all of which are
chitinous 65. These membranes become separated
from the general surface of the gut, but tend to remain
attached at the anterior end, and are therefore apt
to give a false impression of taking origin solely at
this point. They remain attached in this region
largely because here the mid-gut is overlaid by the
oesophageal valve. That is probably the raison
d,'etre of this invagination: it enables the peritrophic
membrane to extend well beyond the point at which
the food enters the mid-gut.
Now as the food pa.sses through the oesophageal
valve it will tend to press the walls outwards; and
in so doing it will tend to press the secretion from
the cells of this cardiac region into a membrane. That
is the crudest form of mechanism by which the peri-
troplllb membrane is pressed out; it is met with in
the w,rvae of Lepidoptera 122, 280. But in many
insects the conditions have become far more elaborate.
The !lells in the cardiac region have become increas-
ingly, specialized for the production of the membrane
until, in the Diptera and in the Dermaptera, they alone
are concerned in its formation. At the same time
the mechanisms by which the fluid secretion from
these cells is pressed to form a solid tube become
more complicated, until in many Diptera these
annular moulds or presses present the most elegant
forms 280. Sometimes the oesophageal valve bears a
rigid cuticular ring against which the cells of the mid-
gut are forced (Fig. 8, A, C); sometimes the valve is
itself a solid structure made up of large vesicular
cells (Fig. 8, B) ; sometimes, a rather cruder arrange-
ment, the valve is thin-walled but contains blood
sinuses which can be distended with fluid and so
blown out against· the cardiac cells.
The peritrophic membrane is present in the majority
of insects. It has disappeared in the Hemiptera,
' ...
i,. .. --pm""
c
FIG. S.-Annular moulds producing peritrophic membrane:
A, larva. of mosquito, Anopheles; B, tsetse fly,
GZo88ina ; C, earwig, Forjicula. (Modified after
Wigglesworth.) The cross-section of the membra.ne is
equal to that of the mould in each case; hence the
complicated folding of the membrane in the narrow part
of the mid-gut in B.
cr, cuticular ring fonning inlier wall of press; en" circular muscle coinpress-
Ing the outer wall against this rillg; do, duct of crop; m, sphincter .jIluscle ;
mg, mid-gut; 0, oesophagus; pm, peritrophic membrane; Be, cells secreting
the substance of the membrane.
perhaps because these take only liquid food, and in

various blood-sucking forms such as the Culicidae,
Tabanidae, fleas and sucking lice; it is wanting also
in the Carabidae and Dytiscidae, possibly because
their digestion is largely extra-intestinal (see p. 50),
possibly bec{tuse their secretion is of the holocrine
type (see p. 50) in which the secreting cells break.
DIGESTION 49
down completely during the process. It is absent
also from adult ants 85.
Salivary Glands. Most insects possess salivary
glands. Sometimes, as in the Apterygota 198 and in
the bee 159, several kinds of glands, probably with
very different functions, open in the region of the
mouth; and even the single pair of labial glands
present in most insects have very varied functions.
(i) In the first place, their secretion may serve to
moisten and dissolve the food: when a cockroach
eats, its mouthparts are regularly moistened with
saliva 278 and in the feeding butterfly a drop of saliva
is extruded from the tip of the tongue 243. (ii) Then
their secretion often contains digestive enzymes
which may act outside the body (as in the plant-
sucking Aphids whose saliva dissolves the starch in
the plant cells) 19 and which continue to act upon
the food after it has been swallowed (amylase in the
cockroach 247, invertase in butterflies 243 and bees 150).
(iii) In the Hemiptera, Diptera and Siphonaptera,
which have mouthparts adapted for sucking up fluids,
the saliva is discharged at the tip of the proboscis,
and one of its functions may be to keep the tube, up
which the fluids are sucked, moist and clean. (iv) And
in those forms which suck the blood of living animals,
the saliva usually contains an anticoagulin-probably
antithrombin 58. It has been shown that if the
salivary glands are removed from the living tsetse fiy,
although it continues to take food for some time,
ultimately its crop and proboscis become blocked
with coagulated blood 176.
There is, however, much still to be learned about
the salivary glands of insects. In many Hemiptera
they consist of several lobes, which differ in histo-
logical structure and in the staining reactions of their
secretion. What the functional significance of these
differences may be is at present quite unknown.
It is interesting to note that many predaceous
beetles (larvae of .Dyti8CU8, Oarabu8 and Oieindela,
and the adults of these forms) which lack salivary
glands, as well as the ectoparasitic larvae of certain
Hymenoptera, eject their intestinal secretion from
the mouth, allow much of the digestion of their prey
to take place outside the body, and then reabsorb
the fluid prodqcts. This process is termed extra·
intestinal digestion 174,. The flesh-eating maggots
achieve the same result by discharging proteolytic
enzymes in their excrement 127.
Secretion. The secretion of digestive juices in
insects is commonly stated to be of two histological
types: holocrine and merocrine. In the former type,
which occurs notably in Orthoptera and Coleoptera,
the active epithelial cells disintegrate complet~ly in
the process of secretion, and are replaced by the
growth of new cells from special cell nests or nidi 234.
In the merocrine type, the cells do not break down.
They may discharge their secretion, usually through
a striated border, without any visible changes in the
cell (save what may be detected by refined cytological
methods) and that is probably the most common
state of affairs. But a special type of merocrine
secretion, known as vesicular secretion, is described,
in which discrete droplets of fluid are eliminated
through the striated border 44. Unfortunately, iden-
tical droplets are only too easily produced artificially
during fixation, and appear in abundance where
during life they cannot be seen at all. But this does
not, of course, exclude the possibility that in some
cases vesicular secretion may actually take place.
Sometimes the cells of the gut contain conspicuous
vacuoles, and these are often taken as evidence of
secretory activity. But it must not be forgotten
that such vacuoles may equally well arise in the
course of absorption "46. In Aphides, fed after a period
of fasting, successive waves of secretory activity
appear to pass down the stomach 268.
The Reaction of the Gut. As was discovered
long ago by Plateau, the intestinal contents of insects
DIGESTION 51
are not as a rule strongly acid or alkaline. The saliva
in those forms in which it has been examined is usually
neutral 276, 277; the gut contents usually slightly
acid, having a pH between 6 and 7 183. The contents
of the gut tend, on the whole, to be more alkaline
in phytophagous insects (pH 8·4-9·0 in some herbiv-
orous Coleoptera, pH 8·4-10·3 in the silkworm 235)
and more acid in carnivores 19; but there are many
exc~tions 163. Well-marked acidity (pH 4·8-5·2) may
develop in the crop of the cockroach, after a meal
of carbohydrates, as the result of acid fermentation
by micro-organisms 276; and sometimes the insect
itself is responsible fOl'a quite strongly acid secretion;
as in a well-defined region in the intestine of the blow-
fly maggot, which has a pH of 3·0-3·5, the acidity
being due probably to phosphoric acid 127.
Digestive Enzymes. The digestive enzymes of
insects are such as might be predicted from the nature
of their food. Omnivorous forms, like the cockroach,
have a full complement of enzymes capable of digest-
ing all the common foodstuffs. 24 7 Insects which take
chiefly a protein diet, such as the exclusively blood-
sucking species, have little but proteolytic enzymes. 277
Where an insect feeds only on nectar, as in the adults
of Lepidoptera, only an invertase is present 243 ;
whereas the phytophagous larvae of these same forms
have a protease, lipase, amylase, maltase and inver.
tase 243. A few examples to illustrate this general
relation between the nature of the food and the
enzymes present, are given in Table I; many more
such exaJIlples could be quoted. It will be seen that
most of the enzymes are secreted in the mid-gut;
the salivary glands rarely secrete anything but
amylase, and often contain no enzymes at all.
These enzymes are readily demonstrated in extracts
from the gut. They are present also in the juice
removed from the lumen 247, showing that at least
the greater part of digestion is extracellular. Some-
times attempts have been made to demonstrate the
existence of enzymes by observing the time of survival
of insects upon pure foodstuffs 259. Such experi-
ments may throw valuable light upon the significance
of particular substances in nutrition 284, but they
cannot, of course, prove the existence of enzymes;
for positive results may be due to absorption without
hydrolysis, or to bacterial fermentation.
The properties of the enzymes of insects are, in
general, very like those of the corresponding enzymes
from vertebrates 278. They are similarly affected by
changes in hydrogen ion concentration, and are stim-
ulated or inhibited by the same factors. The chief
proteolytic enzyme seems always to be more or less
like pancreatic trypsin, * though it often acts best over
a rather more acid range of pH 127, 277. Enzymes of
the pepsin type acting in very acid media do not
occur. Certain special enzymes are present in those
insects which feed on particular substances; for in-
stance, the meat-eating maggot (LuciZia) has a
collagenase which attacks raw connective tissue in
alkaline medium 127; and a few of the wood-boring
beetles produce a cellulase 219. (The clothes moth
(Tineola) can digest keratin, and the bee moth
(Galleria) can digest wax-in both cases probably
without the aid of bacteria-but active preparations
of the enzymes concerned have not been obtained 183.)
So many insects feed on the foliage and woody
tissues of plants that particular interest attaches to
the question of cellulose digestion. In the vast
majority of species, the cellulose is quite unaffected
by passage through the intestinal canal: those plant
cells that are not ruptured by the jaws of the cater-
pillar pass through the gut with their walls intact,
although their contents may be digested 20; the
excrement of many wood-boring beetles contains all
the lignin and cellulose that have been ingested with
* The proteolytio enzymes of inseots have not yet been
studied on modern lines with the object of differentiating
their trypsin into its oonstituent elements.
DIGESTION 53
TABLE I
Enzym68
FoOli
Salivary
Glanda Mid-gut
OBTHOl'TERA
Cookroach (Peri- Omnivorous Amylase Amylase; maltase;
l'laneta ameri- invertase; lactase
cafIG) ... (in very small
amounts) ; tcyp-
tase; peptidase;
lipase.
COLEOPTERA
Japanese beetle Foliage Maltase; invertase ;
(PopiUiajapon- tryptase; lipase.
ica)·"
HYMENOPTIt~
Honey-bee (Apia Nectar and Amylase; Amylase; invertase;
meUijlca) II, "8 pollen invertase tryptase; lipase.
LEPIDOl'TERA
Silkworm (Bom- Foliage - Amylase; maltase;
byz mon) invertase (chiefly
Larva ... in the cells); tcyp-
tase ; peptidase
(only in the cells);
lipase (in very
small amounts).
Humming bird Nectar Invertase Invertase.
hawk moth
(MacroglosBtJ
eteUatarum)
Adult •••
DIl'TERA
Blowfly (Lucilia Flesh Amylase Tcyptase; collagen-
B6ricata) (very ase ; peptidase ;
Larva II. weak) lipase.
Blowfly (OaUi- Liquid food Amylase Amylase; maltase;
l'hora) ofaIIkinds invertase; weak
Adult ••• tryptase and pep-
tidase.
OhryBOpB rilacea Female: Nil. Very weak amylase;
Adult 181 blood. strong invertase;
Male: nec- tryptase ; pepti-
tar, &c. dase.
Tsetse-fly (Gl08- Blood Nil. Very weak amylase;
sifIG)m strong tryptase ;
pe~tidase.
the food 163; the very starch grains may be protecte~
from digestion by their pectin covering. On the
other hand, many insects can digest c~llulose; some-
times by means of cellulase they secrete themselves 1119,
sometimes with the help of symbiotic micro-organisms.
Symbionts in Digestion. Many insects have a
rich fauna of bacteria Or protozoa in the gut, but in
none is this so prominent as in the hind-gut of wood-
boring termites. Here there is an amazing population
of flagellates, ciliates and spirochaetes, which perform
for their host the invaluable function of digesting
cellulose 66. Termites with this fauna intact can
thrive indefinitely upon pure cellulose; but if the
insect is deprived of these organisms (by exposing
it to a high tension of oxygen, for example) its powers
of digesting cellulose are completely lost, and unless
reinfected it soon dies 56. Cellulase can be extracted
from the flagellates in the gut of these termites and
of certain cockroaches (Oryptocercus); whereas this
enzyme is absent from defaunated insects 1161.
Certain Lamellicorn beetle larvae (Oetonia, OrycteB.
Osmoderma) that feed on pine-needles and such like,
ingest with their food those micro-organisms which
ordinarily ferment cellulose in nature. These flourish
in the dilated hind intestine (the so-called 'fermen-
tation chamber ') of these larvae, and form an impor-
tant, perhaps an essential, aid to digestion 27°_being
themselves digested later by the proteolytic enzymes
of their host 1173. The same occurs in some Tipulid
larvae 46. But there are related beetles (Dorcus)
which have a similar fauna and yet are said not to
digest cellulose 219. Other insects that feed on wood
harbour yeast-like organisms, often within the cells
of the gut; and it was natural to attribute to these,
also, the function of digesting cellulose. But, in fact,
some beetles that have such' symbionts' are unable
to digest cellulose, and others which are without them
can do so 163. Many blood-sucking insects, also,
possess intracellular bacteria or yeasts in their tissues ;
DIGESTION 55
and the occurrence of these in close association with
the gut, in the case of the tsetse-fly (Glossina) and the
Pupipara, led to the suggestion that they were con-
cerned in the digestion of blood 48. On the face of
it this was not a probable hypothesis, for the blood
proteins do not differ specially from those of other
tissues; and it has been shown (at least in the tsetse-
fly) to be incorrect; for no digestion of the blood
takes place in that anterior part of the mid-gut in
which the symbionts occur 277. If they are true
symbionts, these organisms must have some other
function (p. 72).
Absorption. In the cockroach, in which the food
is digested largely in the crop, a small amount of
absorption, notably of fats, may also occur in that
segment of the fore-gut 1; but the greater part of
absorption undoubtedly takes place in the mid-gut.
The cells seem never to take up solid particles; the
dissolved foodstuffs, like the digestive enzymes,
diffuse through the peritrophic membrane, when such
is present, and are absorbed by the epithelial cells.
In the blood-sucking insects, such as the tsetse-fly
or the mosquito, almost nothing is passed on to the
hind-gut, save a little haematin 284. In these forms
the hind-gut clearly plays no part in absorbing food-
stuffs; but whether this is generally true is uncertain.
We have seen that the greater part of digestion in
wood-eating termites and Lamellicorns takes place
in the hind-gut; to what extent the products are
absorbed here, or returned to the mid-gut, is not
known; but the hind-gut of Lamellicorn larvae has
special areas of cells which are believed to be con-
cerned with absorption 273. Water is certainly
absorbed in the hind-gut and rectum of many insects
(p. 65); and the food residue, in contact with the
rectal glands, may be converted to a more or less
dry faecal pellet 284; but whether other substances,
also, are absorbed has not been satisfactorily demon-
strated 243.
CHAPTER V
EXCRETION
T HE function of the excretory organs of animals

is to maintain a more or less constant ' internal
environment'. To this end they may be called upon
to eliminate several classes of substances: (i) sub-
stances, like mineral salts or water, which are present
in simple excess in the diet; (ii) the end products
in the metabolism of organic nitrogen, sulphur and
phosphorus, of which, particularly of nitrogen, there
is always a large surplus for excretion-these form
perhaps the most important category of excretory
substances; (iii) more or less complex compounds,
not uncommonly pigmented, which arise perhaps as
accidents or by-products in the course of other
chemical changes, and which haye to be excreted
_because the chemical equipment of the body is power-
less to- deal with them; (iv) and, finally, in order to
maintain such properties of the blood as the hydrogen
ion concentration, or the osmotic pressure, the excre-
tory organs may be called upon to remove certain
substances such as acids or bases, or to hold back
others such as water.
Excretion of Dyes. Organs may fairly be re-
garded as excretory if they perform these functions,
or eliminate one or more of these classes of substances.
But, in practice, it is not always easy to recognize
these activities; and therefore a second, experimental,
criterion of excretory organs has come to be used-
the excretion of dyes introduced into the body. This
test must be used with discretion; for different dyes
56
EXCRETION 57
are absorbed by different tissues: methylene blue,
for example, is taken up by the nervous system, by
the tracheal cells and by the oenocytes, and is there-
fore of no use for this purpose 133. The most reliable
dye is indigocarmine, which seems to be removed
specifically by the excretory organs of all animals.
When indigocarmine is injected into insects, it rapidly
appears in the Malpighian tubes, which are un-
questionably excretory organs 78. In the primitive
Apterygota it appears, also, in the lower segment or
labyrinth of the labial glands 42, and it is probably
correct to regard these also as excretory organs,
homologous perhaps with the excretory glands of
Crustacea. That is all the information obtainable
with indigocarmine.
But if the acid dye ammonia carmine {which, in
vertebrates, is filtered through the glomerulus of the
renal tubule) is injected into insects, it is not excreted
by the Malpighian tubes, but is absorbed and segre-
gated by special cells-notably, by the 'pericardial
cells' which lie along the dorsal heart and aliform
muscles 62, by the' garland-like strand' discovered
by Weismann in the Muscid larvae, by various
scattered groups of cells 43, and, in the Apterygota,
by the upper segment or saccule of the labial gland 42.
On this evidence Kowalevsky put forward the view
that these cells are excretory organs (so-called ' acid
excretory organs', because litmus is turned red within
them) analogous in function to the kidney glomerulus.
, Nephrocytes.' Kowalevsky's experiments were
so striking that this theory at once gained acceptance;
but it is doubtful if it will bear close examination.
Our ideas on the kidney glomerulus have' changed
so much in the last forty years that Kowalevsky's
analogy carries no weight at all. If the ammonia
carmine cells are really 'storage kidneys' (nephro-
cytes, athrocytes) as the theory implies, the waste
substance which they accumulate during life should
at least be visible. But in many insects nothing
seems to accumulate within them at all 62; 138. Often
they contain pigmented granules 136; but this is the
case with many cells-in the fat body, epidermis,
tracheal matrix and so forth. Occasionally, they
may become laden with some pigment from the
food 147; and it cannot be denied that they are then
serving as 'storage kidneys' for this particular sub-
stance; but this can scarcely be their chief function.
Perhaps they play some undetermined part in inter-
mediary metabolism 136; or they may be exoretory
organs in the sense of synthesizing the waste products
that are to be eliminated by the Malpighian tubes 62.
They are highly capricious in their absorptive pro-
perties: they will take up various proteins; they
will absorb some dyes, but not others; and it has
not been possible to recognize any common feature
in molecular structure or physical or electrical pro-
perties to account for these differences 138. But these
properties are very doubtful evidence of an excretory
function; for, in vertebrates, similar properties are
shown by the reticulo-endothelial system and by the
connective tissue fibroblasts.
Urate Cells. As was observed by Milne Edwards,
and later by. Fabre, there are cells in the insect body
which become laden with crystalline spheres of uric
acid (' urate cells '); and they surely can be regarded
as storage ..kidneys, for uric acid is an undoubted
excretory substance. In many cases this view is
probably correct; throughout larval life in the social
Hymenoptera, uric acid collects in solid form in
special urate cells scattered throughout the fat
body 229, and is not transferred to the Malpighian
tubes (of the adult) until the end of pupal life ; in
LepiBma, Dermaptera and many Orthoptera, this
state of affairs persists throughout life, and the Mal-
pighian tubes discharge very little uric acid 28£. But
even here the observations must be interpreted with
caution. For uric acid is the end product of protein
katabolism; and when protein breakdown is occur-
EXCRETION 59
ring very actively within a cell, it is likely that. the
rate of formation of uric acid may, on occasion, exceed
the rapidity with which it can diffuse from the cell
and be carried away by the blood; particularly if
conditions in the cell become unduly acid. The uric
acid will then crystallize out, and its subsequent
solution and removal will take place very slowly.
That indeed is what appears to happen in the cells
of the fat body in Muscidae during metamorphosis :
granules of uric acid (' pseudonuclei ') crystallize out
within the ' albuminoid' deposits, and not until late
in pupal life is the uric acid transferred to the Malpi-
ghian tubes 208. The same thing happens in Lepidop-
tera; and here the uric acid must be endogenous,
for it is not increased by the artificial introduction
of preformed uric acid 134.
It is probable that not a few of the urate cells of
insects are of this nature---not true excretory organs
but active cells in which uric acid has crystallized out,
as it were, by accident. For instance, the epidermal
cells of some Hemiptera contain uric acid; but it
has been shown that in one case at least, that of
Rhodniu8 prolixu8, the deposition of this uric acid
takes place only at one stage of the moulting cycle
-just before the new cuticle is laid down; that is,
when the cells in question are most active in pro-
ducing chitin-perhaps from the proteins of the
food 286.
Pigments. Now besides this uric acid, there is
in the epidermal cells of Rhodniu8 a red pigment,
which appears at the same time as the uric acid, and
then slowly diminishes 286. Perhaps this substance,
also, is a by-product of the synthetic activity of these
cells; and perhaps that is the origin of many of the
pigments of insects-accidents in metabolism, which
only secondarily acquire their biological significance.
Certainly this is the case with the pigments of the
Pieridae; as was shown by Hopkins 130 many years
ago, these are composed of uric acid and complex
derivatives of uric acid 274 which, instead of being
transferred from the fat body to the Malpighian tubes,
are deposited in the scales of the wings 275. The
same applies perhaps to the red pigment of Vanessid
butterflies; though this is not related chemically to
uric acid 191.
Verson's Glands. It has been observed that
when the silkworm casts its skin, its body may be
powdered with crystals of oxalate and uric acid
derived from the moulting fluid; and this led to the
suggestion that, at the time of moulting, the Malpi-
ghian tubes are thrown temporarily out of action,
and their excretory function taken over by the
moulting glands (Verson's glands) 263. But this idea
has been shown to be mistaken; the excretory sub-
stances in the moulting fluid really come from the
Malpighian tubes, escape through the anus- beneath
the old cuticle, and so spread over the surface of the
body 233. (It is just possible that the calcareous
warts of Stratiomyid larvae (p. 3) may also be
formed in this way. On the other hand, the lime
which appears in the ecdysial fluid of certain Diptera
at the time of pupation is said to be dissolved from
the Malpighian tubes, reabsorbed into the blood, and
excreted through the newly-formed cuticle 146.)
The Malpighian Tubes and the Urine. When
all these subsidiary processes have been considered,
tlie Malpighian tubes still remain unquestionably the
chief excretory organs. The Malpighian tubes are
relatively simple tubular glands which open at the
junction between the mid-gut and the hind-gut.
They are exceedingly variable in form 269: some-
times being numerous (e.g. over 100) and short, some-
times fmv in number (e.g_ 2) and long; sometimes
simple and sometimes branched; occasionally an-
astomosing to form closed loops; while sometimes
more than one type of tube is present 78. Their his-
tological structure is no less vatiable. Usually their
epithelial cells bear a striated border, but this may
EXCRETION 61
be wanting; sometimes this border is of the type
known as 'Wabensaum' in which the rod-like
elements are fused together; sometimes it is made
up of separate mobile filaments (' Burstensaum ' 283).
Sometimes the tubes have a muscular coat and are
capable of active movements; more often they show
only slow twisting movemepts due to changes in the
secretory pressure within them 78. Many histological
changes have been described in the active cells of the
Malpighian tubes; the discharge of vesicles, the
eruption of vacuoles, and so forth. It is not improb-
able that more than one cytological mechanism of
excretion may exist; but certainly many of the
recorded observations are artefacts 283.
The urine which the Malpighian tubes secrete is
equally varied. It may be a clear fluid or a thick
pasty suspension: the colourless fluid passed by
blood-sucking insects soon after a meal is an example
of the former sort, the' meconium' of newly emerged
Lepidoptera of the latter. Its constituents depend
naturally upon the nature of the food, but it always
consists primarily of water with the usual inorganic
salts, chlorides and phosphates of sodium, potassium,
calcium and magnesium, in solution. Yellowish or
greenish pigments (' entonio-urochrome ') are often
present; but it is likely that these differ chemically
in different species 78. Nitrogen is excreted chiefly
as uric acid.; but urea has been reported in substantial
amounts in the urine of the clothes moth 283, and
meat-eating maggots are said to eliminate much of
their nitrogen as ammonia 12? The amino acid
leucine has been recorded by many earlier authors,
but the evidence rests only on crystal structure and
requires chemical confirmation 78. Guanine, the main
nitrogenous constituent in the urine of many Arthro-
pods, seems not to occur in insects; but the nitrogen
compounds of insect urine are still very incompletely
studied. The reaction of the urine naturally varies
with the diet, and the period after feeding·; the thick
meconium of Lepidoptera has a pH of 5.8-6.3 120 ;
the similar urine of the bug Rkodnius proZixus, shows
the same kind of range 283.
When uric acid is present in the urine it is usually
in the form of crystalline spheres with a radial stria-
tion. These spheres are commonly stated to consist
of ammonium acid urate, or of sodium or potassium
acid urate; but in Rhodnius, almost the only insect
in which this question has been investigated chemi-
cally, they consist mainly of free uric acid 283; and
it is possible that this is also the case in other insects.
Beside these crystals of uric acid, the Malpighian
tubes may contain solid spheres or granules of
calcium or magnesium carbonate 75, 205 and, notably
in the larvae of Lepidoptera, crystals of calcium
oxalate 78. The oxalate is possibly derived from the
preformed oxalic acid in the food, but the significance
of the accumulated carbonates is uncertain. They
are sometimes used by the pupating larva to streng-
then its cocoon (Cerambycidae, Col.) or its puparium
(Agromyzidae, Dipt. 1(6), or by the female to calcify
the shells of her eggs (Phasmidae, Orth. 205), but these
!J.re probably secondary adaptations. It has been
suggested that the carbonates may constitute a
method of eliminating carbon dioxide 146; but such
a mechanism seeJJlS uncalled for, and in any case the
amount of carbon dioxide bound in this way must
form but a fraction of what the insect produces.
Perhaps these carbonates simply provide a mechanism
for getting rid of excess alkali; it is at least note-
worthy that they occur chiefly in saprophagous and
phytophagous larvae of Diptera (Stratiomyiidae.
Drosophilidae, Agromyzidae and many more) whose
food might be expected to contain an excess of fixed
base. Their urine must therefore be alkaline, and
in the presence of the carbon dioxide of the tissues,
if calcium or magnesium ions are present in quantity,
carbonates will tend to precipitate. It is worth
noting that carbonates are present in the phytQ.
EXCRETION 63
phagous Agromyzid Acidia, but not in the parasitic
genus Oryptochaetum 251; and they are said to be
absent in aquatic larvae from very acid waters 165.
(It may be noted in passing that similar deposits of
lime (' calcospherites ') may occur in the fat body of
some insects 148.)
Excretion in Rhodnius. Now insects are essen-
tially terrestrial animals. Consequently, the main
problem with which their excretory system is faced
is the elimination of all these substances, and par-
ticularly the elimination of nitrogen, in such a way
that they are able to conserve their limited supplies
of water. And it is instructive to consider the ex-
cretory system from this point of view. It will be
convenient to start by describing the excretory
system of the blood-sucking bug Rkodniu8 prolixus,
which has been studied in considerable detail 263, and
then to compare with this some other insects.
The excretory system of Rkodnius consists of four
very long Malpighian tubes, opening into a large
rectal sac (Fig. 9, A). Each tube is made up of two
distinct segments: a translucent upper segment,
about two-thirds of the :whole, the lumen of which
contains only a clear fluid, and an opaque white lower
segment stuffed with spheres of uric acid. The histo-
logical structure of these two segments is quite
different; .the one changes abruptly into the other;
and immediately below the junction the granules of
uric acid appear in the lumen (Fig. 9, B). The ex-
planation of these facts which has been suggested is
that the upper segment of the Malpighian tube is
secreting a solution of sodium or potassium acid
urate, and the lower segment reabsorbing both the
water and a large proportion of the base (sodium or
potassium-perhaps in the form of bicarbonate)
leading to a precipitation of the free uric acid. For
it can be shown (i) that the contents of the upper
segment (pH = 7·2) are definitely more alkaline than
those of the lower segment (PH = 6·6); (ii) that
neutral red added to the blood of the insect is secreted
into the lumen by the cells of the upper segment and
reabsorbed again by the cells of the lower segment;
and (iii) that if two ligatures are applied to the tube
near its lower end at a time when the uric acid spheres
have been washed out by a recent meal, then uric acid
FIG. 9.-A, excretory system of RhodniuB: schematic;

B, detail of Malpighio.n tubes at junction of upper and
lower segments; arrows indicate the circulation of water
and base; C, two ligatures applied to Malpighian tube
three hours after meal; D, the same 24 hours later.
(Modified after Wigglesworth.)
Is, lower segment of Malpighlan tube: lumen full of uric acid spheres ;
mg, mld·gut; r, rectum; rg, rectal gland; UI, upper segment of tube.
appears above the upper ligature and below * the

lower ligature, but between the ligatures there is no
uric acid, nor is there any distension of the tube
(Fig. 9, C, D).
Thus it is suggested that, in Rhodnius, there is So
• This has doubtless been derived from the other Mal-
pighio.n tubes, and has entered from below.
EXCRETION 65
continuous circulation through the excretory system
both of water and of base, the same water and base
being used over and over again to carry uric acid
from the body.
It is not unlikely that the rectum, also, assists in
the process of reabsorbing water; for around the
point of entry of the Malpighian tubes there is a ring
of large epithelial cells constituting a so-called rectal
gland; and we shall find that there is evidence in
other insects that the function of the rectal glands
is the absorption of water from the excrement.
Excretion in Other Insects. This conception
of' a circulation of water through the excretory
system is a familiar one in the physiology of birds,
and reptiles, and mammals, where the renal tubule
itself, the cloaca, and the large bowel all take part in
the reabsorption of water. It is an idea that has
only recently been introduced into the physiology of
insects 284; but it helps to explain many of the pro-
perties of their excretory system.
The simplest system that occurs is that shown in
Fig. 10, A. Here the Malpighian tubes are composed
of a single type of cell, and contain only fluid. As
this fluid, mixed with the intestinal contents, passes
down the 4ind-gut, water is absorbed from it-first
by the cubical epithelium of the intestine and finally
by the epithelium of the rectum. The material is
retained in the rectum until it has been converted
into a more or less dry pellet. In some cases, for
instance in the human louse Pediculu8, the contents
of the mid-gut may be retained in the stomach so
that the hind-gut contains only urine; under these
circumstances, a little pellet of solid uric acid may
be formed in the region of the rectal glands, clearly
demonstrating the absorption of water by these
organs 284. This arrangement is found in Lepi8ma,
Dermaptera, Orthoptera, Neuroptera, and many
beetles. In some cases the active epithelial cells of
the rectum are collected into compact areas, the
'rectal glands'; in other cases they are evenly
spread over the gut wall.
In many beetles, phytophagous, carnivorous and
omnivorous species 183, as well as those that feed on,
FIG. lO.-Types of excretory system in insects: schematic

Hlnd·gut indicated by a heavy line Internally. Explanation in text.
dry substances 210, such as the mealworm (Tenehrio),

the upper parts of the Malpighian tubes closely invest
the rectum, being bound to it by a delicate membrane
(Fig. 10, B). The precise significance of this arrange-
EXCRETION 67
ment is not known; perhaps it serves to add the
absorptive powers of the Malpigruan tubes to those
of the rectal epithelium; but the arrangement seems
always to be associated with remarkable powers of
drying the excrement. The rectum of the mealworm,
for instance, contains a bone-dry powder, consisting
largely of uric acid, while the Malpighian tubes secrete
only a clear fluid 284.
A similar arrangement is met with in the larvae
of Lepidoptera (Fig. ]0, C) 143. But here the lower
part of each Malpighian tube contains solid granules
of uric acid, oxalate, &c. 78; and this suggests that,
as in Rhodniu8, the Malpighian tubes themselves assist
in reabsorbing water.
In many Diptera (the mosquito, Muscid flies, &c.),
the Malpighian tubes may contain solid uric acid
throughout their length (Flg. 10, D) 284. The tubes
consist apparently of but a single type of cell, so
that the mechanism for the precipitation of uric acid
must be different from that in Rhodniu8. In the
rectum, a further absorption of water probably occurs ;
but these insects lack the power of really drying the
excrement. Conditions uxe somewhat similar in
certain Hymenoptera. In other Hymenoptera, the
excretory system is like that of the flea .(Fig. 10, E).
It never contains any solid uric acid because sufficient
water is retained in the capacious rectum to keep it
in solution; but it is none the less probable that this
fluid is reabsorbed, and circulates again through the
system 284.
Finally, in those many dipterous larvae in which
the Malpighian tubes contain carbonates, these are
generally confined to the upper part of the tube
(ll'ig. 10, F); but the mechanism of their secretion
is unknown 75, 205.
These are just a few of the arrangements of the
Malpighian tubes and hind-gut, which will serve to
illustrate the general principles of excretion in insects
so far as these are known at present; many other
arrangements exist. And it should be pointed out
that many other hypotheses as to the function of the
rectal glands of insects have been put forward from
time to time 284. Some of these are certainly in-
correct; but others are probably subsidiary functions
in particular species. _
Excretion in Aquatic Insects. Lastly, it is
interesting to consider what changes take place in the
excretory system of insects when they come to live
in fresh water, and are no longer compelled to be so
careful of their water supply. This matter has re-
ceived little attention; but in the case of the mosquito
larva, there is around the anus a group of hollow
papillae, theso·called' analgills'. These are covered
with chitin, but with chitin devoid of its cuticulin
coat and therefore, unlike the rest of the body surface,
freely permeable to water. In superficial appearance
these organs look strikingly like rectal glands that
have prolapsed through the anus; and indeed it has
been shown that they also are engaged in absorbing
water, not from the excrement but from the environ-
ment 280. This water is excreted by the Malpighian
tubes, which are kept so well flushed out that normally
they never contain any solid uric acid. But if the
larvae are reared in isotonic or slightly hypertonic
salt water, in which absorption no longer tak~s
place, masses of solid granules frequently make
their appearance in the lumen of the Malpighian
tubes 280.
Accessory Functions of the Malpighian Tubes.
Reaumur observed that the lemon yellow powder
used by the larva of M alacosoma neustria to cover its
cocoon is a product of the Malpighian tubes. Such
products are used by other Lepidoptera: Leucoma
salicis, Eriogaster lanestris and so forth 64; and we
have already seen how Cerambycidae strengthen their
cells with lime from this same source. The foam or
cuckoo-spit pJ;:oduced by the larvae of Cercopidae is
formed from the Malpighian excretion mixed with
EXCRETION 69
the product of some other glands 288. And in the
larvae of such widely separated insects as Lebia
(Carabidae) and certain Neuroptera, a portion of the
Malpighian tube is transformed into a silk gland 84.
CHAPTER VI
NUTRITION AND METABOLISM
INCE the alimentary canal of insects contains

S digestive enzymes competent to hydrolyse those
constituents commonly present in their food (p. 51),
it is probable that this food is absorbed in the form
of simple products of hydrolysis: carbohydrates as
monosaccharides, fats as glycerol and fatty acids,
protein as amino acids. But at present there is no
proof of this, and it is possible that there may be
some absorption of more complex bodies. Little is
known, either, of the fate of the absorbed products.
Presumably, the tissue cells take from the blood what
they require for growth, energy production or secre-
tion; and surplus food is certainly taken up by the
cells of the fat body, in which it is stored as glycogen,
fat and protein. But what is quite unknown is where
the intermediary stages of metabolism take place.
IJ amino acids are condensed to form protein before
storage, where is this done? Where is carbohydrate
converted into fat? When unwanted proteins or
amino acids are deaminated, is this effected by thosc
tissue cells which require the products, or by some
organ specially devoted to this branch of metabolism ?
It is pOSSible, though no very certain evidence of
this yet exists, that the pericardial cells may be respon-
sible for some of these -activities (p. 57) 136; and it
has been suggested that the oenocytes may be con-
cerned either in some general process of intermediary
metabolism 133, 160, or in the special process of syn-
thesizing certain constituents of the cuticle 286. On
70
NUTRITION AND METABOLISM 71
the other hand, the accumulation of solid uric
acid in the fat body when protein reserves are broken
down (p. 58), and in certain cells of the cuticle during
moulting (p. 59), suggests that the powers of deami-
nation and uric acid formation are shared by many
tissues; though the chemical details of both these
processes are unknown.
Food Requirements. We have seen how exceed-
ingly varied are the apparent food materials of insects
(p. 42). But it must be· remembered that insects
are not always feeding upon what they eat: they
may devour rotten wood but feed only upon the fungi
which it harbours; or eat large quantities of cellulose
and yet assimilate only the associated substances 259.
The principles of insect nutrition are therefore not
easy to state. It is self-evident, though it is some-
times forgotten by writers on the subject, that for
growth and reproduction, insects must have an
adequate supply of carbon, and nitrogen, phosphorus
and sulphur, inorganic salts and all the other elements
that go to make up the living system. There is, at
present, little information as to what may be the
simplest substances which will satisfy these needs in
a given insect; but there is certainly no reason to
suppose that they are any simpler than those required
by vertebrates. Drosophila has been reared with
ammonium salts 259, and cockroaches with glycine 296
as the sole source of nitrogen; but these insects have
in the intestine a notoriously rich flora of yeasts and
bacteria which may well have been responsible for
essential syntheses. For the maintenance of life and
activity, on the other hand, without growth, insects
can subsist for long periods on carbohydrates alone,298,
and Lepidoptera usually live only on sugars through-
out their adult life. The hexose monosaccharides
which can or cannot be metabolized under these
circumstances are much the same as· those in other
animals, but (for the bee at any rate)' the pentoses
seem to have no food value 284. There are much
wider differences in respect to the di- and tri-sacchar-
ides, due no doubt to the presende or absence of
appropriate enzymes 264.
Vitamins and Symbiotic Organisms. One
aspect of nutrition that has been explored consider-
ably is that relating to accessory substances or
vitamins 259, and the part played by micro-organisms
in providing these, The larvae of the blowfly Lucilia
can be grown from sterile eggs on the sterile brain of
mammals; they will not grow on certain types of
sterile muscle, But if the muscle is infected with
suitable micro-organisms, or if a sterile extract of
yeast is added to it, normal growth is obtained, The
bacteria and the yeast provide a vitamin (belonging
to the B group of vitamins) that is present in brain
but lacking in muscle 128, It is interesting to note
that sterile blood, also, is an insufficient diet for these
larvae; it can be rendered adequate by the addition
of vitamin B I 128, Now many insects (the bed-bug
(Oimex), the sucking lice, the tsetse-fly (Gl0881,na), the
Pupipara) take no food throughout their life except
sterile blood, But all these insects * contain within
their bodies special groups of cells (mycetomes) stuffed
with supposedly symbiotic micro-organisms 46, which
are absent from such insects as mosquitoes, fleas, &c"
that take blood during part only of their life cycle.
This naturally suggests that these symbionts may
constitute an endogenous source of vitamin, and thus
enable these insects to grow on sterile blood 8, 277;
indeed, it has been shown experimentally that if the
louse Pediculu8 is deprived of its mycetome and
symbionts, its powers of nutrition and reproduction
are greatly impaired, and that this impairment may
be made good, to some extent'at least, by the addition
of yeast extract to the diet 7,
'" The blood-sucking Reduviidae (RhodniUB, Triatoma, &c.)
are an exception, and have no intracellular symbionts. But
they regularly have the gut contents heavily infected with a
diphtheroid bacillus or 8 streptothrix, the source of which
has not yet been determined SBB,
Fat soluble accessory substances, more or less
related to vitamin A, are perhaps also required by
insects 217, 246; vitamin C seems not to be neces-
sary 246; but some insects apparently need factors
differing from those known in vertebrate nutrition 128.
It is clear, therefore, that when insects feed upon a
sterile and restricted diet, the supply of vitamins may
become a problem of some urgency; and it is possible
that the association of sterile food (as in the majority
of plant-sucking Hemiptera) with the presence of
symbionts, may again be traced to this need. But at
present there is no experimental proof of this, and
difficulties are presented by the occurrence of sym-
bionts in the omnivorous cockroach, and their presence
in some wood-boring insects (p. 54) but not in all 46.
The same difficulty, of course, is encountered in any
of the suggested hypotheses of symbiotic function:
whether the symbionts are digesting cellulose 46,
fixing atmospheric nitrogen 117, or synthesizing vita-
mins, it is not easy to see why related forms (Ceram-
bycid larvae, for example), with and without
symbionts, should grow with equal rapidity on the
same diet 219. But the insect and its symbionts show
such wonderful mutual adaptati!)ns, that it is equally
difficult to dismiss the whole phenomenon as one of
harmless parasitism.
Wat~r Metabolism. Another fundamental re-
quirement in insect nutrition, and a very important
one, is water. Even such an insect as the weevil
Calandra, which is accustomed to feed in dry materials
and is highly adapted to conserve the water in its
body, cannot survive unless its food contains some
10 per cent. of water 142. In other words, the water
produced from the oxidation of foodstuffs, although
of great importance to the insect, is not, alone, suffi-
cient for its needs. If the mealworm is kept 'in air
with a relative humidity of 90 or 95 per cent., it is
able, by some mechanism that is quite unexplained,
to absorb water from the atmosphere 185; but this
property is very unusual; most insects cannot absorb
water vapour even when the air is saturated 52, 67.
On the other hand, the loss of water (when this occurs
by evaporation as opposed to excretion (p. 56)) is
markedly affected by the water vapour in the air.
The rate of loss, like that from inanimate bodies, has
been found, within certain limits, to be more or less
proportional to the saturation deficiency of the air 62.
It is not, of course, to be expected that such a law
should hold very exactly, especially at varying tem-
peratures, because evaporation occurs chiefly from
the tracheal system 109, 188, the ventilation of which
varies with the intensity of metabolism in the insect
and so with the external temperature (p. 21) 115.
But apart from the importance of water as a con-
stituent of the tissues, it plays a significant part in
the reaction of the insect to extremes of temperature
in its environment. When exposed to high temper-
atures, an insect may be killed (depending upon the
conditions of experiment) by the temperature itself
or by the desiccation consequent upon evaporation 186.
Now, provided the insect in question is sufficiently
large, it will be able to lower its temperature below
the fatal level (for short periods at any rate) by
evaporating water from its body, and thus to exercise
a limited degree of temperature control 186. And it
has been found that insects which have lost a certain
_amount of water by desiccation are, for some un-
known reason, more resistant to high temperatures:
normal individuals of Leptinotarsa die at 58°-60° O. ;
desiccated ones at 1°-5° higher os. The same loss
of water also makes them more resistant to low tem-
peratures 142: if the larva of the beetle Popillia
japonica is experimentally dehydrated to half its body
weight, it becomes exceedingly resistant to cold, and
can survive exposure to -28° 0. 207 • In association
perhaps with this phenomenon, it is usual for insects
to reduce their water content before going into hiber-
nation; this reduction is certainly one factor in
the increased resistance of insects to cold during the
winter 207. When the water content is reduced in
this way, the freezing point and undercooling point
(the lower limit of supercooling) of the whole insect
are lowered-partly perhaps because the crystalloids
in the tissue fluids are increased in concentration,
but partly because a greater proportion of the water
is influenced by its association with the body colloids.
H it be assumed that water which remains unfrozen
at _20 0 C. is 'bound' to the colloids, then it may be
said that there is an increase in ' bound' water during
hibernation. In winter· hardy species, this increase
may take place without any change in the total water
present 220.
It seems clear that the relation of water to the
tissue colloids is one of the factors in cold hardiness;
but there is so little agreement among colloid chemists
on the subject of ' bound' and' free ' water that, at
the present time, any generally acceptable explana-
tion of the facts in terms of colloid chemistry is not
possible. It has been suggested, also, that the binding
of water to tissu.e colloids is one factor in the resistance
of insects to desiccation 220.
The Temperature of Insects. We have seen
that at high temperatures, insects above a certain
size can keep themselves cool for short periods by
evaporation of water from the body, or by making
use of evaporating water in the environment 187.
At low temperatures, their bodies are often kept
warmer than the surrounding air by the chemical
changes going on within them. And both these
methods of temperature control are used by the social
Hymenoptera in maintaining the temperature of their
nests or hives 126. It seems that for the intense
muscular activity of flight, a high body temperature
is necessary: the large Sphingid moths are apparently
unable to take flight immediately from rest; they
first stand with the wings vibrating-shivering, as it
were-until the temperature in the thorax has risen
above 30° C. Only then can they fly; and during
flight the temperature will exceed 40° C.72. Similar
changes occur in other insects during activity··280;
but whether insects at rest will increase their meta-
bolism in order to maintain the body temperature is
at present uncertain.
Besides generating heat by muscular activity,
many insects make use of the sun's rays to warm
their bodies to the optimum temperature 164, 260,
their behaviour varying with the temperature
attained; and it has often been suggested that the
tendency of insects in cold climates to be dark in
colour, is related to the absorption of radiant heat
from the sun-though it is sometimes forgotten that
this same dark colour will also favour the loss of heat
during the night 222. The range of external tem-
peratures over which insects are normally active
shows remarkable differences in different species, as
does their resistance to high temperatures 260. But
the physiologioal basis of these differenoes is not
known.
Respiratory Metabolism. The energy meta-
bolism of insects varies, naturally, with the degree
of musoular exertion. It therefore varies enormously
in different species, and is muoh greater in the larva
than in the pupa, and greater again in the adult 161.
It depends also upon the temperature; but a rise in
temperature not only increases the resting or ' basal'
metabolism, but usually leads to greater aotivity.
Only in very exoeptional circumstances, therefore, is
any simple mathematioal relationship likely to be
found between these two factors. Curves of various
types, relating temperature with oxygen uptake, not-
ably during pupal development, have been described
mathematically by some authors; while others regard
such curves as being really composed of a number of
straight lines whioh intersect at 'critical points'
where new metabolio processes are supposed to super-
vene 260.
The energy metabolism of such an insect as the
bee, when at rest, is of the same order of magnitude
as that of man: about 20 gm. calories per kilo. per
minute. But during flight 145, this may be increased
nearly 1,300 times, and the bee weighing 100 mg.
may then burn sugar at the rate of 35 to 40 mg. per
hour 158_:whereas in man, doing the maximum
amount of work possible, the respiratory metabolism
is increased only some 10 or 12 times. During
activity, the respiratory quotient approximates to
unity. During starvation, the quotient falls-in-
dicating that, as in other animals, the stores of
carbohydrate are consumed first, to be followed by
protein and fat 57, 181.
Anaerobic Metabolism. Insects are remarkably
resistant to lack of oxygen. Down to a low level of
oxygen tension, the precise level varying in different
insects, the oxygen uptake remains unchanged 114, 125.
Below this level it falls off; but many species can
survive for long periods in the complete absence of
air. The extra metabolism of activity very quickly
ceases, and the insect becomes quiescent; the basal
metabolism continues anaerobically. The type of
chemical change does not alter: the unoxidized meta-
bolites simply accumulate. Thus, by the time the
insect is again restored to the air, it is greatly in
arrears for oxygen; and in order to payoff this
, debt', and oxidize the lactic acid and other sub-
stances that have accumulated, its rate of oxygen
consumption remains above the normal level for a
long time (Fig. 11, B) lI36. During the anaerobic
period, the blood naturally becomes more acid, the
pH falling, in the case of the grasshopper, from 6·8
to 5'8, and the carbon dioxide capacity is reduced liD";
consequently, at very low tensions of oxygen, there
may be a liberation of combined carbon dioxide 114.
This type of temporary anaerobIc metabolism is a
normal occurrence in the life of Gastrophilus (para-
sitic in the stomach of the horse) which can survive
as long as 17 days in the complete absence of
oxygen 71; experimentally, it occurs in insects of all
kinds.
Now it has been repeatedly observed that during
the pupal stage of insects, the rate of oxygen con-
sumption first falls, and then rises again as the
imaginal organs are formed; giving, altogether, a
characteristic U-shaped curve 2011. The commonest
explanation for this curve is that the intensity of
Ol~--~~------~~
A
days
FIG. n.-Ordinates: ra.te of oxygen uptake; a.bscissae:
time
A, oxygen uptake during the whole pupal stage of 91 days in Galhria, Lep.
(after Taylor and Steinbach); B, oxygen uptake in grasshopper, MeltJnopllU.
During the 75 minutes that the curve coincides with the base line, the insect
was submerged in water (after Bodine).
metabolism is a measure of the amount of formed

tissue present--the larval tissues in the early period,
the adult tissues at the end. But another possible
explanation, which does not seem to have been put
forward, is that during the reconstruction of the
tracheal system, metabolis_m becomes largely anaero-
bic, and the insect accumulates a temporary oxygen
debt, of the SmIle kind-though of course less in degree
-as in the insect in nitrogen. There are three obser-
vations which support this notion: (i) during the
pupal stage, the blood often becomes more acid and
the carbon dioxide capacity falls 22; (ii) very low
respiratory qUQtients are often observed, indicating
an incomplete combustion of fats 179; and (iii) a
definite fall in oxygen uptake may be observed at the
end of the pupal stage-suggesting that the oxygen
debt has been paid"off (Fig. 11, A) 249. It is of interest
to note that during both the pupal stage (in Tenoorio
and other insects) 260 and the egg stage (in Aciliu8,
Col.) 161, the total oxygen consumption during
development is the same at all temperatures, although
the rate of development varies enormously. Thus it
would appear that the energy requirements of develop-
ment are independent of the temperature.
Chemical Changes during Metamorphosis.
The changes in the total composition of the insect
body during metamorphosis naturally vary in different
groups, with the different types of morphological
change 202. As a rule, the large stores of glycogen
accumulated towards the end of larval life (these may
exceed 33.per cent. of the dry weight of the body in
the six-day-old bee larva 237) fall steadily, almost
reaching zero in the developed adult. The fat
decreases rapidly at first, then more slowly (during
the period of depressed oxygen consumption) and
then rapidly again before emergence. Fat does not
usually fall so low as glycogen, but only a small per-
centage of the original store persists in the adult.
In the pupa of the Muscid Lucilia, it appears to be
the unsaturated fatty acids which are consumed, the
saturated acids remaining more or less unchanged 84.
The protein is sometimes largely used in the formation
of the cocoon (the silkworm uses ha1f its body protein
for this purpose 202); the remainder is extensively
broken down during the early part of pupal life, the
non.protein nitrogen, particularly the amino acids,
increasing. With the growth of the new tissues, the
process is reversed, and the amino acids fall 69 (p. 37).
The inorganic phosphate follows the same course as
the amino acids 59.
These changes, and the corresponding changes in
other c'onstituents and in the respiratory quotient,
represent, of course, only the summation of the
processes of synthesis and breakdown that must be
going on throughout metamorphosis. At times it
must happen that these two processes will exactly
balance in respect to a particular substance over a
given period, and then chemical analysis will reveal
no change at all. It is clear, therefore, that such
analyses of the whole animal are of rather limited
interest; or rather, are impossible to interpret in-
tellJgently, until they have been correlated with the
histological sequence of events. Conversely, intricate
cytoplasmic changes are described in the fat body
and blood cells during metamorphosis, the chemical
significance of which is almost completely unknown ;
though it appears as though nuclear substance, fat
and protein were being merged into a single material
easily transported to and assimilated by the growing
imaginal tissues. 21
Suspended Metabolism. From time. to time,
certain of the metabolic activities of insects become
suspended for more or less prolonged periods. These
periods of rest or, as it is termed, diapause, may
supervene at any stage: egg, larva, pupa or adult.
They are characterized by a reduction of activity
in the motile stages, but more particularly by the
cessation of growth-arrested development in the
egg, larva and pupa, failure of the reproductive organs
to mature in the adult. Therefore, although the
respiratory metabolism is reduced during diapause,
and, when it occurs in the pupal stage, the period of
low oxygen uptake is enormously prolonged 118, dia-
pause is perhaps best regarded as a problem of growth.
In many insects this arrest may be brought about
by a variety of adverse conditions-cold, drought,
inadequate food and so forth-and normal growth
is at once restored when conditions again become
normal 60 ; or dormancy may be a direct effect of
temporary desiccation 52. In other insects the arrest
is not brought to an end in this way, and diapause
continues until broken by some other stimulus,
although the environmental conditions have again
become favourable. And finally, there are many
insects, in regions with a well-defined cold or dry
season, which regularly spend the unfavourable
months in a state of diapause;, in them, this state is
assumed as the result of an inborn rhythm which
manifests itself even when external conditions are
uniformly favourable 80,63.
Various hypotheses have been advanced to explain
the occurrence of diapause: that it is an auto-
intoxication from accumulated waste products 221,
that it is the expression of an hereditary rhythm in
the life cycle, and so forth. But the essential feature
of diapause seems to be the cessation of growth. We
shall see that growth is probably maintained by
internal secretions (p. 93). The most probable hypo-
thesis in regard to diapause is, therefore;thatitis due
to a failure in the hormones necessary for growth 287,
induced in some cases by nutrition or by environ-
mental conditions, in others by an inborn rhythm.
The varied stimuli which will' break' the diapause:
pricking, burning, severe cold 221, the oviposition or
development of parasites 261, &c., may be pictured as
evoking again the production of these hormones.
An alternative hypot,hesis would picture diapause as
due rather to a second factor which temporarily in-
hibits the normal growth process 30.
Pigment Metabolism. We have already seen
that in the oourse of metabolism pigmented sub-
stances may arise as waste products (p. 59). Pig-
ments may also be absorbed :with the food and aocu-
mulate within the body: the flavone pigment in the
wing-soales of the butterfly Melanargia 142, for
example; and the carotinoids in the blood of the
potato beetle Leptinotarsa, and in the epidermal cells
of the bug Perillus which preys upon it 160. But it
is remarkable that apparently similar pigments in
more or less related insects may be quite unrelated
chemically: thus, the common red and orange pig-
ments of many Hemiptera may be carotinoids from
the food, anthocyanin-like or flavone-like pigments,
perhaps also from the food 121, 150, or pigments of
unknown nature formed in metabolism 286; and the
Pieridae may use their purine colours to mimic other
butterflies the chemistry of whose pigments is. quite
different 130. Similarly with the green pigments: in
many cases these are synthesized by the insect itself,
and have no relation to the chlorophyll of plants,
even in phytophagous species; for their development
is unaffected (in Dixippus 102, in locusts 86, and in
many caterpillars 191) by rearing them upon food con-
taining no pigment; nor have they the chemical
properties of chlorophyll and xanthophyll 86, 191.
The sexual differences in the blood of many cater-
pillars, which have been ascribed to differences in the
utilization of plant pigments, are probably due rather
to differences in the composition of the haemolymph
proteins 191. On the other hand, it is impossible to
dogmatize on this subject; there are many who still
believe that much of the green pigmentation of insects
is due to altered chlorophyll 1411, 282, and even that
genetic differences in colour may result from differ-
ential absorption of the pigmented elements of which
chlorophyll is composed 100.
The other type of insect pigment is the insoluble
black substance melanin, which is formed in the
presence of oxygen by the action of an oxidizing
enzyme upon some colourless aromatic precursor,
the' chromogen'. Melanin-producing enzymes are
generally diffused throughout the blood and tissues
and secretions of insects, particularly near the time
of pupation when much black pigment is often
formed 70; the chromogens are usually deposited in
restricted areas, and so determine the colour patterns
that develop 107.
The melanogenic enzymes can act upon a great
variety of chromogens: tyrosine (monoxyphenyl-
alanine), 'dopa' (dioxyphenylalanine) and various
other polyphenols; and the enzyme concerned is
ttlrmed tyrosinase, 'dopa' -oxidase, polyphenolase,
according to the substrate used. But it is not certain
whether in fact more than one enzyme is present, and
in few cases has the natural chromogen been recog-
nized. In the tissues and in the cocoons of certain
Lepidoptera (Ramia cecropia, Eriogaster lanestris,
&C. 212 ) and in the elytra of Melolonth~ 228, the spon-
taneously oxidizable ' dopa ' has been demonstrated;
and in the mealworm Tenebrio, dioxyphenylacetic
acid has been proved to be one of the chromogens
present 228. But it is, of course, very probable that
more than one chromogen may be concerned in a
given insect; indeed, one view of the significance of
melanin formation is that it provides an excretory
mechanism for disposing of sundry toxic phenols
arising in metabolism. Experimentally, the quantity
of melanin formed can be widely varied by changes
in the external conditions, notably the temper-
ature 121, 160, 227. In SOlJle cases (Habrobracon,
Hym.), these effects may he transmitted to the off-
spring up to the second generation 14.4.
Melanin may be incorporated in the substance of
the cuticle (pp. 2, 11), or deposited in granules in the
epidermal cells. The other pigments may be in
solution in the blood or in the fat droplets of the cells
(lipochromes), or deposited as granules in the epider-
mis or other tissues, or between the layers of the wing
scales.
In some insects the coloration may be strikingly
influenced by external factors. Sometimes, notably
in the stick insect Dixippus, such changes are rever-
sible-the so-called 'physiological colour change' .
There are no discrete chromatophotes in these cases,
there is simply a migration of pigment granules, a
clumping or spreading, in all the epidermal cells lOS.
In Dixippus this change may be brought about by
a great variety of stimuli: temperature, narcotics,
osmotic changes in the blood, mechanical pressure,
all of which act directly upon the epidel'Itlal cells;
and optical stimuli and humidity, which act through
the eyes and tracheal system respectively, and then,
by way of a centre in the brain, lead apparently to
the secretion of a hormone into the blood, which
determines the pigment movements 8, 102. These
pigment changes follow, also, a diurnal rhythm (the
insect is dark by night and pale by day) which, though
determined primarily by illumination, may persist
for several weeks in complete darkness 102.
In these same insects, if the factors or stimuli
already enumerated continue to act over long periods,
the amounts of the different pigments formed may
be permanently changed; and this may be effected,
also, by nutrition, particularly by the moisture of the
food. This permanent effect is termed 'morpho-
logical colour change'; but when brought about by
optical stimuli, at any rate, it appears again to be
produced, like the temporary change, by way of the
nerve centre and the internal secretion 8. The co-
ordination of these mechanisms may lead to remark-
able resemblances to their background in various
Orthoptera 86.
As was shown by Poulton many years ago, larvae
and pupae of certain Lepidoptera also possess won-
derful powers of acquiring the tone or coloration of
their background. This phenomenon has been
analysed most carefully in the case of Pieris pupae.
In these the quantity of pigment deposited in the
cuticle appears to be determined by the specific
quality of the light received into the eyes of the
pupating larva, acting through a centre situated in
the head 35.
Another very striking colour change is that in
locusts, the solitary phases of which are quite differ-
ently pigmented from the gregarious phases. The
gregarious type of coloration can be evoked by
crowding the young forms together, and there is
evidence that the factor responsible for the colour
change is the incessant muscular activity which over·
crowding causes. It has been suggested that the
active muscles produce a substance (provisionally
called 'locustine ') responsible for this change 86 ;
but it is perhaps worth considering whether the factor
in question may not be an increase in the acidity of
the blood resulting from the ceaseless muscular con-
traction, acting perhaps directly, perhaps by way of
the pigmentation centre.
CHAPTER VII
REPRODUCTION AND GROWTH
T HE reproduction of insects forms one of the

most remarkable chapters in their natural
history. The structural adaptations for copulation,
the control of the sperm in the body of the female,
the micropylar arrangements in the egg, which allow
fertilization through the fully formed shell, are re-
markable enough. But the curiosities of insect re-
production: parthenogenesis, occurring sporadically,
regularly or, in some species, apparently exclusively;
viviparity, sometimes of newly hatched larvae, some-
times of single full grown larvae that have been
nourished in the uterine cavity by special 'milk
glands'; polyembryony in parasitic insects, in which
a single egg may give rise to several hundreds of
larvae; and paedogenesis, where the ova in the young
ovaries of larvae or pupae ripen prematurely and
develop into larvae whic1} disrupt their parent host-
all these phenomena present most fascinating prob-
lems 141. Sometimes these abnormal paedogenetic
or parthenogenetic generations may alternate with
generations of the ordinary sexual type; and it is
this alternation, due sometimes to inborn factors,
sometimes, apparently, to environmental conditions,
which make these matters specially interesting from
a physiological point of view. But, at present, the
physiological factors at work are quite obscure, and
in this chapter we shall confine our attention to the
more prosaic aspects of subsequent growth.
Growth. The phenomena of growth and develop-
86
REPRODUCTION AND GROWTH 87
ment in insects are, in broad outline, like those of
higher animals. At one pole of the egg there is a
centre (analogous, perhaps, with the organization
centre of vertebrates) which determines differentiation
in the embryonic germ 216, 231; once differentiation
has begun, development; co-ordinated by some un-
known mechanism, proceeds until the insect hatches;
then follows a period of growth, encUng in maturity
and reproduction. As in other animals, there is often
a change in the proportions of the body as it increases
in size; and when this is the case, the disproportion
follows the law of disharmony or heterogeny.* This
law, the physiological significance of which is quite
unknown, applies equally to morphological dimen-
sions and to the concentration of chemical con-
stituents 250.
But upon this general background are superimposed
some special phenomena, conditioned by the make-
up of the insect; and it is in these that we are mainly
interested. In the first place, the cuticle is so con-
structed that once formed it cannot grow in surfa.ce
area. If it is soft and much folded it may Btretch
gradually, and then the insect may increase Con-
tinuously in length as it does in weight. This happens
in the larvae of flies (e.g. DroBophila) and Lepidoptera
(e.g. Galleria) 250; but even in:these insects there are
hard regions of the cuticle which can grow only by
replacement at the time of moulting. The growth of
such parts is, therefore, discontinuous; and when
the greater part of the insect is encased in a rigid
cuticle, the growth in length of the whole body is
likewise discontinuous, and takes place in a series of
steps (SphodromantiB 214, DixippUB 80, GerriB 250).
It was observed long ago by Dyar, working with
the larvae of Lepidoptera, that the linear dimensions
of the hard parts increased at each Il!oult by a constant
* That is, that the logaritlun of the dimeIl!!ion of the part
is proportional to the logarithm of the dimension of the
whole (y = KX").
7
ratio, usually of the order of 1·4 (Dyar's law) 141.
More recently, it has been shown that various non-
metabolic insects (Sphodromantis 214, Dixippus 80)
approximately double in weight from one moult to
the next, while their linear dimensions increase 1·26
(i.e. iY2) times. This growth factor of 2 was first
demonstrated by Przibram and Megu~ar 214; it is
perhaps best regarded simply as a special case of
Dyar's law 250; but it is sometimes referred to as
'Przibram's factor', and an attempt has been made,
first by Przibram himself and later by others, to
prove that this factor is universally applicable to
insect growth 26. This is only possible by introducing
the conception of 'latent divisions' and supposing
that the insect can multiply its weight by 2, perhaps
several times, without moulting. In this way the
growth of any insect can be made more or less to
obey the same law. The physiological basis proposed
by Przibram to explain his factor was that at each
moult (or at each latent division) every cell in the
insect body divides into two. But what histological
evidence exists is definitely opposed to this improbable
hypothesis. For in the Muscidae, almost the whole
of lantal growth is accomplished by increase in size
of the cells without multiplication 208, and in hemi-
metabolic insects there is much cellular breakdown
and reconstruction' besides the irregularly distributed
cell divisions 286.
We have seen that at the time of moulting, the
insect enlarges its surface area by swallowing air
or water (p. 13). In the latter case (Ae8chna 232,
Notonecta 250, &c.) the resulting curve for growth in
weight is peculiar and shows an abrupt rise immed-
iately after moulting, followed by a more or less level
period during which the ingested water is gradually
incorporated as true growth (Fig. 12, B). Equally
curious is the growth curve of the blood-sucking bugs
(Oimex, Rhodnius) which take only a single gigantic
meal during each moulting stage, so that the weight
curve shows a series of acute peaks gradually rising
to a higher level (Fig. 12, C) 51.
Moulting. The chief interest of insect growth
centres round the phenomenon of moulting or
ecdysis 148. This is essentially a process of growth ;
for it is the only means by which the hard parts can
.,.
Ime
FIG. 12.-Growth curves; semischematic: A, Di:cippU8,
(modified after Teissier); B, GerriB, Hem. (modified
after Teissier); C, Rhodniu8, Hem. (modified after
Buxton)
The &rrOwslndicate the tlme of moultlng; the Insect becomes adult at ~
increase in size (p. 12). ThE)re is no doubt, therefore,

that growth is one factor that determines moulting.
But in some insects (Tineola 256, Tenebrio 250) moulting
can occur repeatedly during starvation and without
any enlargement: Tineola may' so~etimes develop
a sort of 'moulting fever' and cast the skin as many
as 'eight times in two or three days 268; and adult
Collembola will continue to moult without change in
size or organization 250. There must therefore be
some other factor at work. It is natural to suppose
that this factor is of the nature of a hormone; and
there is some experimental evidence of this, for if
the blood of normal caterpillars is injected into other
caterpillars near their time of moulting, the onset of
moulting is delayed; but if the blood of moulting
caterpillars is injected, moulting is not delayed. From
which it is inferred that the process has actually been
accelerated 48. More convincing evidence is afforded
by the fact that epidermal structures transplanted
from one caterpillar to another of different age, even
of a different species) undergo ecdysis at the same
time as their new host 27. And the presence of a moult-
ing hormone has been clearly demonstrated in some
recent experiments with the bug Rhodniu8 prolixu8 287.
Rhodniu8 requires only a single feed in each of its
five stages; moulting takes place at a definite interval
after each meal. In the fifth or final nymph, this
interval is about 28 days. Now, if the recently fed
insects are decapitated before a certain 'critical
period' (about 7 days after feeding), they fail to
moult, although some of them may survive more than
twelve months. If decapitated later than this, they
will moult. But by allowing the blood from an insect
decapitated 8 days after feeding to flow into an insect
decapitated 1 day after feeding, the latter may be
caused to moult.
The next question is the source of this hormone.
One suggestion is that it comes from the ' moulting
glands' (Verson's glands) in the epidermis 48, 139;
another, that it comes from the oenocytes 153-both
of which organs show a cycle of secretory activity
which synchronizes with moulting 286. But it has been
shown that in Rhodniu8 neither of these hypotheses
is tenable; because moulting begins before the
moulting glands are differentiated and before the
oenocytes reach the height of their activity 288. Now
we have seen that decapitation inhibits moulting in
Rhodnius; which suggests (though of course it does
not prove) that thEl source of the hormone is in the
head. Of the organs in the head, the only one which
shows distinct changes during this period is the corpus
allatum, the cells of which swell up to a maximum
at about the seventh day after feeding, and then
diminish. It seems likely, therefore (although the
evidence is still very incomplete) that the corpora
allata (which are nests of glandular -cells budded off
from the epidermis 201, like the endocrine organs of
vertebrates, and innervated by the visceral nervous
system) are the source of the moulting hormone 287.
As to the stimulus which causes secretion of this
hormone: there is evidence that the stimulus to
moulting in Rhodniu8 is the stretching of the abdomen
by the meal; and section of the nerve cord in the
prothorax inhibits moulting like decapitation. This
suggests that moulting is started in Rhodnius by a
nervous stimulus to the brain. But this cannot, of
course, be a general explanation, applicable to those
insects which moult repeatedly though starved.
Metamorphosis. Growth in holometabolic in-
sects (Coleoptera, Lepidoptera, Hymenoptera,
Diptera), that is, insects in which the adult is mor-
phologically very unlike the young, and in which a
resting pupal stage is interposed between the final
larva ann the imago, is generally pictured as a com-
petition between farval and imaginal cells. Through-
out larval life the "former have the upper hand, and
the imaginal cells, in the form of single' histoblasts '
or clusters of undifferentiated cells termed ' imaginal
germs', are held in an embryonic state. Metamor-
phosis takes place when the larval cells die (' his-
tolysis ') and the imaginal cells escape from the
influence that has restrained them, and proceed to
grow and differentiate (' histogenesis '). That is, of
course, a very crude conception of the process; for
there is no absolute distinction between larval a.nd
imaginal tissues. Even in the extreme case of the
Diptera, many larval organs (Malpighian tubes;
certain muscle groups) are remodelled without much
change to form thos~ of the adult 208; and, con·
versely, a certain amount of histolysis takes place
even during the ordinary larval moults 211, 286.
Thus, physiologically, metamorphosis differs from
moulting only in degree: it is a moulting process,
typically occupying two stages, in which the morpho.
logical changes are extreme. Now at the final moult
of hemimetabolic insects (Hemiptera, Odonata, &c.)
there is, again, a morphological transformation which
is far more extreme than at the preceding moults.
It is probably justifiable, therefore, to compare the
physiology of this final moult of these' exopterygote '
insects with that of metamorphosis in the 'endo·
pterygote' groups; and to suggest that the two types
of development are likely to be controlled by similar
factors.
Many hypotheses have been advanced to explain
the initiation of metamorphosis: the intoxication of
the larval tissues by accumulated waste products 141 ;
their asphyxiation by carbon dioxide 141, or their
autolysis induced by other acids 22; the failure of
hypothetical secretions from the larval cells, which
are supposed to have held the embryonic adult tissues
quiescent, and kept the phagocytes at bay 208; the
overgrowth in size of the larval cells resulting in their
death from starvation 254 ; the awakening of the adult
tissues by secretions from the genital glands 208, or
by tyrosinase secreted into the blood 70.
The current conception is that metamorphosis, like
moulting, is initiated by a hormone, and that thiS
hormone differs in quality from that which causes
moulting 27. We have seen that decapitation of
Rlwdniu8 nymphs, before a certain critical period
after feeding, prevents moulting; after this period,
normal moulting occurs. But if the fourth., third.,
second., even the first.stage nymphs are decapitated
around the critical period, a certain number of them
suffer a premature 'metamorphosis' and develop
the characters of diminutive adults 287. One must,
therefore, conclude that the head exerts two in-
fluences: (i) it initiates moulttng; (ii) it inhibits the
realization of the imaginal characters until the insect
is fully grown. And.if one may generalize from a
single hemimetabolic insect, and apply this inter-
pretation to holometabolic insects, then the most
probable mechanism of the control of metamorphosis
is the inhibition of the imaginal germs, and of imaginal
differentiation in general, by a hormone controlled by
the brain. This hypothesis has already been put
forward by Kopec to explain metamorphosis in
Lymantria, but upon very indirect evidence which
he himself has later come to doubt 157. On the other
hand, Kopec has produced good evidence that the
brain (or some closely associated organ) is necessary
for the initiation of the pupal moult 167.
This conception, of two competing hormones, one
inducing growth, the other inhibiting metamorphosis,
provides an explanation of 'prothetely '-the occa-
sional appearance of pupal characters in mature larvae
of certain beetles (Tenebrio, &C. 175) and Lepidoptera
(Lymantria, &C. 104)-if it be supposed that this phe-
nomenon represents a partial escape of the embryonic
tissues from the restraining influence of the i:nhibitory
hormone. This condition finds a parallel in Rkodniu8 ;
for of those early nymphs decapitated around the
, critical period', while some show unmistakable adult
characters, others show only some slight modifications
towards the adult form 287.
Further, the condition of Rhodnius nymphs de-
capitated before the 'critical period', which slowly
digest and metabolize their food, but fail to grow, is
very suggestive of the state of diapause in normal
insects (p. 80).
Histolysis and Histogenesis. There has been
much controversy as to the mechanism of breakdown
of the larval tissues at metamorphosis. All that is
certain is tha.t at a given stage the imaginal histoblasts
first begin to proliferate rapidly; then each tissue
that is to suffer histolysis disintegrates and dis-
solves 211. It is uncertain whether the cytolytic
agents come from the dying tissues themselves or
whether they are secreted into the blood I by the
leucocytes or other cells 208. Sometimes the phago-
cytic cells in the blood invade and seem by their own
activities to disrupt apparently healthy tissues 208 ;
as a rule, obvious cytolysis is visible before the arriving
phagocytes absorb and digest the products 208; some-
times the whole process of solution takes place without
the intervention of phagocytic cells 203, 1137; and often
all these types of breakdown occur in di:(ferent parts
of the same insect 254. The final result is the same:
the products of cytolysis of the larval cells serve to
nourish the growing tissues of the adult; though the
intermediate stages of the process, which mliSt
obviously differ from one group of insects to another,
are not clearly known.
The co-ordination of all these processes to produce
the adult form is the problem of orderly growth which
is at present quite unsolved. The potentialities of
form seem generally to be inherent in the tissues
themselves (wings, eyes, integument, &C. 155) and to
be more or less independent of the other organs in
~he body. Though the development of the muscles
in metamorphosis seems to depend upon their con-
nexion with the central nervous system 157.
Regeneration. The general obscurity of the
factors controlling growth is well illustrated by the
phenomena of regeneration. In the adult insect there
are some slight powers of hea.ling of wounds 173 with
the formation of new cuticle, but no extensive powers
of regeneration. In the pupa the same is true; if
the developing wings in the pupa of Tenebrio are cut
short, they round themselves off and approximate
to the normal form, but there is no true regeneration
of tissue 182. If the wing germs of Lepidoptera are
mutilated in the last larval stage, the damaged areas
of pattern are not made good; whereas if this is
done in the preceding instar, wings of normal propor-
tions but reduced size r~generate; and if the germ is
divided, reduplicated wings result 182. At a still earlier
stage, the wing germs can be extirpated completely
and are formed anew from the hypodermis 182.
Clearly, throughout development, as differentiation
proceeds, there is a progressive loss of totipotence and
regenerative power 213; in the pupa everything is
finally determined except for relatively slight changes
in colour pattern (p. 98) 87. In hemimetabolic insects,
if the appendages are removed in the early larval
stages, they show progressive regeneration at suc-
ceeding moults, and may attain almost normal pro-
portions by the time the insect becomes adult. Here
again there is a progressive loss in regenerative power
as the insect grows older, which is due partly, but not
entirely, to the reduction in the number of moults
in the older insects, and the corresponding reduction
in the opportunities for growth 213.
In all these cases, the removal or partial removal
of an organ results in more or less accelerated growth
at the site of the wound. In the case of a limb, the
new growth is most rapid if removal is effected at the
level where autotomy (the spontaneous separation
of an appendage) occurs naturally; but it is not
necessarily connected with the property of autotomy,
for there is regeneration if appendages are cut through
above or below this level, and there is regeneration
in appendages which cannot autotomize 245_
The influence which moulds the outward form of
the new appendage seems to reside in the epidermis
itself: regeneration is unaffected by extirpation of
the corresponding ganglion; though when this is done,
neither muscles nor nerves are formed within the
new appendage 156, 245. In some cases, regeneration
is 'beterQ:~Il(:)q>bQUB': in DixippU8, if the antenna
is cut through at t~e level of the flagellum, it regener-
ates as an antenna; if it is amputated through one
of the two basal segments it commonly regenerates
as a leg, complete with tarsal claws lI4. But though
this tendency exists, there is no absolute relation
between the level of section and the type of regenera-
tion 98.
Thus, the phenomena of regeneration, like those of
normal growth, can be defined by certain laws-but
they are laws of growth and of regeneration; they
cannot yet be expressed in the simpler terms of phy-
siology, far less in terms of physics and chemistry.
Determination of Sex. The genetic theory of
sex determination has been admirably described by
Dr. F. A. E. Crew in this series of monographs 81.
This theory, which is based very largely on studies
of insects, will explain the determination of sex both
in ordinary sexual reproduction and in partheno-
genesis. It can probably be accepted that, in insects,
both primary and secondary sexual characters are
determined by the genetic make-up of the body cells
-the secondary characters are not evoked by internal
secretions from specia,l glands, either in the gonads
themselves or elsewhere. For (i) removal or trans-
plantation of the gonads at any stage of life is without
effect upon the secondary sexual characters 155 ;
(ii) where the genetic constitution, in respect to sex,
of some body cells differs from that of others, a sexual
mosaic or gynandromorph results, parts of which are
entirely male and other parts entirely female 81 ; and
(iii) if the wing germs of Lepidoptera of one sex are
transplanted into the other sex, their sexual char-
acters are not influenced by their new environment 155.
But in insects, as in other animals, the genetic sex
can be, overridden by other factors, and it is these
other, physiological, factors in which we are inter-
ested here. The phenomena are most conveniently
described in terms of Goldschmidt's well-known theory
of intersexes 105. Leaving aside the genetical analysis,
REPRODUCTION AND GROWTll 97
this theory supposes that sex i& determined by a
quantitative balance between male determining and
female determining factors within the nucleus of
every cell. By analogy with the hormones of the
animal body, these competing influences are often
pictured as being chemical in nature (although, of
course, they might be of some quite Qther kind)-
that is, as hormones working inside the cell.
In the normal female the quantitative influence of
the female determining factor outweighs that of the
male; but the male factor is still there, and under
certain circumstances its effect may become manifest.
Thus, the balance between these factors, though
normally determined according to the laws of genetics,
may (according to those who accept this interJlreta-
tion of the facts) be upset in several ways. (i) By
crossing races in which the sex determining factors
are of unmatched strength. Hence 'intersexes'
result, the morphological characters of which are best
explained by supposing that, owing to a difference in
the rate at which the male and female factors come
to exert their influence, the insect has developed up
to a certain point as one sex and then' switched over'
to complete its development as the other 106. (ii) By
abnormal external temperatures, which are supposed
again to upset the rate at which the two factors work,
in such a way that the latent factor, which would
normally be suppressed, is able to exert its specific
effect, so that ..inter-sexual forms result 106. (iii) By
the action of internal parasites-notably the effect
of Stylops upon certain Hymenoptera 223. This
effect was formerly called ' parasitic castration' ; but
it is more than the mere assumption of a neuter con-
dition, it often involves a partial reversal of sex:
females acquire male characteristics, and vice versa.
In other words, the latent sex factor is again manifest-
ing itself. This result seems to be due to starvation;
for it occurs only in those Hymenoptera which receive
a fixed ration to last the whole of larval life, and not
in those social forms which are fed according to their
individual needs; and the effects are exaggerated if
more than one parasite is present 223. In this con-
nexion it is worth recalling that the sex ratio in certain
Braconid parasites is influenced by the size of the
host in which they mature 131._perhaps this is a
related phenomenon.
Determination of Other Characters. There
is some indication that there may exist a parallel
between the determination of other characters of
bodily structure and those of sex. In other words,
that Goldschmidt's theory may become a general
theory of genetics 106. For the most part, such body
characters are determined genetically; and when
multinucleate eggs are fertilized. by se'Veral sperma-
tozoa carrying different genes, mosaics result in the
adult insect, exactly comparable with the sexual
mosaics or gynandromorphs 108. But characters,
ordinarily hereditable, can sometimes be elicited by
changes in external conditions during development.
These effects can be explained, as the occurrence of
intersexes can be explained, by supposing that many
morphological variations are induced by changes in
the velocities., of particular processes of development.
Such changes in time relations may be brought about,
on the one hand, by changes in the genes, or on the
other, by alterations in the temperature or other
factors, which influence one process more than
another 103, 108.
The importance of rate of development in deter-
mining the final characters is well seen in certain
butterflies with different spring and summer colour
patterns: the summer form (prorsa) of Araschnia
levana is modified into the spring form (levana) by
exposure to cold during a brief susceptible period
following pupation; and the degree of modification
is proportional to the length of time that elapses
before emergence-the slower the individual develop-
meJ}t, the greater is the modification into levana 244.
REPRODtJCTION AND GROWTH 99
It is interesting to note that the coloration centre in
these cases (like the colour centre in Dixippus, &c.
(p. 84)) is in the head: if the head is kept cold and
the abdomen warm, cold forms result 103. Distur-
bances in the velocities of different developmental
processes may explain also the phenomenon of
prothetely 104 (p. 93).
CHAPTER VIII
THE NERVOUS SYSTEM, SENSE ORGANS AND
BEHAVIOUR
NSECTS are endowed with sensory or receptor
Inervous
organs of a high order, linked, through a complex
system, to an equally elaborate effector
apparatus; and the complexity of behaviour that
results from the co-operation of these systems has
yet to be satisfactorily described in physiological
terms. In the future, perhaps, the psychological
method may provide an adequate description of insect
behaviour; but most of the advances of the past
thirty or forty years have been made along the lines
of mechanistic physiology, and it will therefore be
more helpful for our present purpose to approach the
subject from that point of view.
The Nerve Cord. The central nervous system
of insects generally shows a well-marked segmenta-
tion. Typically, each somite has its own pair of nerve
centres or ganglia, giving off sensory and motor nerves,
and connected to the adjacent ganglia by a paired
cord containing only nerve fibres. In virtue of these
ganglia, each body segment enjoys a considerable
degree of autonomy; probably each has its own res-
piratory centre controlling the movement of the
corresponding spiracles, and each is capable of carry-
ing out such reflex movements as do not involve the
activity of other segments. The essential functions
of these ganglia: the nature of the nervous discharge
from the ganglion cells 2, the mechanism by which
certain muscle groups are inhibited while their oppo-
100
THE NERVOUS SYSTEM 101
nents contract 9', even the precise course of the reflex
paths-are problems which have scarcely been studied
in insects.
When a given reflex demands the co-operation of.
a number of body segments, there is situated in one
of the ganglia a higher centre which co-ordinates the
activities of the others. For example, the individual
segments in the larva of Corethra are capable of con-
tracting laterally upon stimulation, but true swim-
ming movements are possible only if controlling
centres in the third and fourth segments of the abdomen
are intact 6; in Cloeon, the respiratory movements
of the gills can be effected by the segmental ganglia ;
but their normal rhythm is controlled by an inhibitory
centre in the second thoracic ganglion and a stimula-
tion centre in the sixth abdominal ganglion 6 ; and so
on for all the movements that have been studied 164.
In those insects in which most movements involve
only the thorax, as in the winged forms, the ganglia
have become enlarged and concentrated, and the
isolated thorax is then capable of performing per-
fectly many of the motor functions-walking, clasp-
ing, vibration of the wings-of which the whole
animal is capable 16'.
The Brain. There is also a concentration of
ganglia in the head: the suboesophageal ganglion
and the supraoesophageal ganglion or • brain'. Of
these, the suboesophageal seems to be the equivalent
of the motor ganglia that we have been discussing;
it controls the movements of the mouth appen-
dages 154, and may contain co-ordinating centres-as
in caterpillars, where it controls the forward walking
movements 154" and in the adult of Aesikna, where
it inhibits the tenacious clasping reflex of the legs 12.
The brain, on the other hand, seems never to contain
motor centres of this kind. It seems rather to be
the central receptor for the abundant sense organs
located in the head; and as such it orders the move-
ments of the whole insect in accordance with the
stimuli whioh these reoeive. The reflex movements
of the isolated body segments are themselves pur-
posive in a limited degree; but when oo-ordinated
by the brain, all the body movements serve the
purpose of the animal as a whole.
If the brain is removed, the reflex reaotions of the
body become exaggerated; it reacts to stimuli which
in the normal insect would have no effect 17; and
movements, such as the cleaning movements in the
bee, may continue without cessation for hours at a
stretch 17. Thus one function of the brain, one
method by which it disoiplines the organism, is to
inhibit the too rigid execution of reflexes by the lower
centres. If one half of the brain is destroyed, the
inseot tends to move in a circle towards the sound
side. This reaction is due partly to the increased
propulsive activity in the limbs on the operated side
following the removal of inhibition 17; partly to an
effect of the brain in increasing the muscular tone
of the flexor musoles on its own side 177. But neither
of these mechanisms will provide a complete explan-
ation of the circus movement; for after mutilation
of the limbs on either side, the performance of the
musoles is so modified that the same movement still
occurs 11. The cause of this behaviour therefore lies
deeper: each half of the brain, when acting alone,
tends to bring about movement towards its own side
~movement that requires the co.operation of muscles
in all parts of the body 4, 12.
Reflex Responses in the Normal Insect.
Comparatively simple reflexes undoubtedly playa
large part in the responses of the normal insect; and
our main prob~em is to describe the whole of insect
behaviour in terms of such reflexes. Pressure upon
the tarsus causes a reflex withdrawal of the leg;
removal of the feet from contact with the ground
evokes a turning over reflex 138 or initiates the vibra-
tion of the wings 110; contamination of the antenna
induces cleaning movements 138. The mechanical
,THE NERVOUS SYSTEM 103
character of such reflexes is well seen in the cock-
roach: if the antenna is stimulated, and the insect
is offered a bristle, it will clean the bristle and neglect
the antenna 138; and many insects will continue the
cleaning movements after their antennae have be~n
amputated. But though mechanical, these reflexes
are purposive and plastic: the water beetle DytisrJUs,
deprived of one hind-leg will swim with the niiddle
leg instead 18; and if the fore-legs of the cockroach
are removed, the stimulated antenna will be held by
the middle pair 138.
Ev'en the simplest of these reflexes involves many
parts of the body, and some movements of the whole
insect are also clearly reflex. This is notably the case
with certain inhibitions. We have seen how loss of
contact between the feet and the ground will initiate
flight; conversely, flight is inhibited when contact
is regained 90. And contact with other parts of the
body may similarly inhibit movement: the apparent
preference for dark crevices in such insects as the
moth Amphipyra 177, the earwig ForjirJUla 271, and the
bed-bug Oimex, depends in fact upon a reflex inhibi-
tion of their movements by contact (' thigmotaxis' or
, stereokinesis '). In the state of immobilization or
, akinesis ' which such conditions induce, these insects
may require strong stimuli to arouse them. Experi-
mentally, a similar hypnotic state may be induced,
in the earwig for example, by many kinds of con-
stantly repeated' stimulation 272; and this state
is doubtless related to the death-feigning reflex
or 'thanatosis' shown by many insects when
alarmed 23, 242.
Then insects moving iLt random may encounter
unfavourable stimuli-too great heat or excessive
cold for instance-whereupon they show a reflex
'avoiding reaction' and turn aside. By this me-
chanism, which is termed' phobotaxis' 166, insects
may be confined within a zone where the stimuli are
optimal: houseflies, Musca, in ~ cold room with a
8
single bunsen burner will collect in a circle at a fixed
distance around this source of heat 260.
Orientation. But neither in 'stereokinesis ' nor
in 'phobotaxis ' is the insect truly orientated; it is
either trapped in a given spot by reflex inhibition, or
restricted to a given zone by , avoiding reactions' 89.
Many insects, however, do show truly orientated
movements (' topotaxis ') in relation to such varied
stimuli as light, sound, gravity, heat and odours 166.
Several mechanisms have been suggested by which
these directed movements might be brought about.
The simplest, known as 'tropotaxis', which is
associated particularly with the name of Loeb, sup-
poses that the tone of the muscles on either side of
the insect's body is determined by the intensity of
stimulus received in bilaterally symmetrical sense
organs 177. That ,being so, the insect is automatieally
turned by its muscles into such a position that stimu-
lation on the two sides is equal; whereupon the
muscula.r power becomes equal, and the insect moves
in a straight line.
Many of the reactions of insects to light -both
simple forms like the maggots of Muscids 177, and
highly specialized animals like the adults of certain
flies 99-can be beautifully explained by this simple
mechanism; and there can be little doubt that it
does playa part in directing the movements' of some
·insects. But it is certainly not the whole story. For
although many insects, when offered two similar
sources oflight, will make for a point between them 170
(as was to be expected if they are directed by the
tone of their muscles), and although many insects
will move in circles if one eye is blackened over (as
though the muscular tone on the two sides of the
body were permanently unbalanced 177), yet after a
time, as the eye becomes adapted to the light, the
higher forms at least will ' learn' to go in a straight
line though they have but a single eye, or, ~th two
eyes, they may select one of two sources of light and
THE NERVOUS SYSTEM 105
go to that 258, And after removal of one or more
legs on either side, they will still make for the light
in a straight line 184,
Such insects seem, in fact, to direct their movement
to a chosen source of stimulus, to the neglect of others,
as though they had the end in view, This type of
orientation is therefore termed' telotaxis '186, The
response seems to be made up of a series of reflexes
specifically related to the region of the eye that is
stimulated 184 (like the fixation reflex in man); so
that the insect finally turns Ulltil the image of the
source of stimulus is received upon the region of
clearest vision (in the insect with a single eye) or
upon the corresponding points in the two retinae (in
insects with both eyes intact) 184, So exact has this
reflex become in such insects as the fire-fly Photinu8,
that the single flash of light emitted by the female
will cause the male to turn accurately in her direc-
tion 184, ,
This same' fixation reflex' doubtless provides the
basis of the response to moving objects that is shown
by many insects, Some forms, like the larva of the
dragon-fly 12, will follow any object moving near
them; others, like the adult dragonfly, after turning
momentarily to such an object, instantly leave it if
it is not suitable for food 12, the initial reflex being
inhibited by the higher centres, In the bug Rhodnius,
the response to moving objects is normally elicited
with difficulty, but it becomes very marked if the
antennae are removed 287,
Then there are insects which will move at a constant
angle to a source of light; a method of orientation
known as 'menotaxis' 186, In some cases this
response seems to result from the balanced turning
effect of two lights-that from the source itself and
that reflected from the background 55, Sometimes
included in this mechanism is the '.~ht-compass'
orientation 50,291 of ants and bees, which make use
of the direction of the sun's rays in finding their way
back to the nest 89. Certain aquatic insects (e.g.
Notonecta) , maintain their position in I1lnning water
by a comparable mechanism-by keeping constant
the image of objects on the bank. If these objects
are moved, the insect can be caused to move with
them 230. Even such insects as Gyrinidae, which
swim in all directions in the water surface, seem, sur-
prisingly enough, to maintain their positibn by the
same mechanism 41.
It will be seen that the present tendency is to
explain many of the reactions of insects to light as
complex reflexes, and to restrict more and more the
application of the muscle tonus hypothesis. The
same is trp.e of the responses to warmth (in blood-
sucking insects) and smell-stimuli which are per-
ceived mainly. by the antennae. It is difficult to
believe that such diffuse stimuli could cause accurate
orientation by differential tonus effects on the two
sides of the body, that is, by 'tropotaxis' 177. It
has sometimes been suggested that odours are located
by 'phobotaxis' (although there is no evidence of
any avoiding reaction to places where there is no
smell) in combination with' tropotaxis ' 199, 267. But
the simplest mechanism is that which postulates a
reflex pursuit of the antennae into the region of
optimal stimulation 189A • This notion brings the
response into line with the reflex response to light,
that is, with' telotaxis '-though it is possible that
at short range, when the stimuli are very .strong,
differential tonus effects may occur. As in the case
of light stimulation, there will be a tendency, as was
noted by Treviranus more than a hundred years ago,
for the insect with a single antenna to follow this,
and so to turn in a circle when exposed to the attrac-
tive scent; and this tendency will naturally be more
marked when the antennae are relatively fixed
structures not capable of individual movement 89.
But none the less, the insect may be able to locate
the source of smell very accurately although it has
SENSE ORGANS 107
only one antenna 199, 287, 289A which it could not do
if its movements were controlled solely by tropotaxis.
Sense Organs: Mechanical Senses. We
must now consider briefly the sense organs of insects
from the point of view of their relation to these forms
of orientation. The guiding sense of many insects
is that of. touch 271, subserved by articulated spines
on the antennae and elsewhere. By administering
suitable punishments and rewards, cockroaches can
be trained, in a maze, to turn always to the right
or the left; but when deprived of their chief organs
of touch by removal of the antennae, this discrimin-
ation is lost 33. The tactile sense is important also
in stereokinesis (p. 103), though exactly what sense
organs are concerned in this response is not estab-
lished.
Another important source of orientation in insects,
notably in the finding offood by many caterpillars 177,
and by the earwig 271, is their movement against
gravity (negative geotaxis). This reaction is not
served by any special statocyst organ, but seems to
depend upon receptors of tension in the muscles or
tendons 14_receptors which must also be of the
greatest importance to the insect in maintaining its
normal posture 14. Closely related to, or perhaps
identical with these organs are the tension receptors
known as chordotonal organs. At their simplest
(i) these serve to detect the external strains and
pressures to which the body is SUbjected 123; (ii)
others are used to detect the strains at the base of
the wings during flight; (iii) there is always a group
at the apex of the pedicel of the antenna (Johnston's
organ) which is probably stimulated by displacement
of the antenna and so serves to detect air currents 77
or, sometimes, even air vibrations or sounds 77; and,
finally, (iv) they are developed ill I!! highly specialized
form, in association with a tympanic membrane, to
provide the auditory organs present in many in-
sects 77, by means of which the females of crickets
and other Orthoptera can recognize and locate the
specific sounds emitted by the males 10.
All these receptors, besides the specific effects which
stimulation of them elicits, have the property of
arousing the nervous system generally, and bringing
it into a reactive condition. They belong to the so-
called 'stimulatory organs' 295 which serve both to
maintain the muscle tone, and to enhance the kinetic
efficiency of the muscles when they are brought into
action. Prominent among the stimulatory organs
are the halteras of Diptera 47 and Strepsiptera 257"",
one important function of which is to sustain the
muscular vigour of the insect during flight by the
stimuli which their own vibration generates. In this
case the receptors seem to be partly the chordotonal
organs but chiefly the abundant campaniform sen-
silla 257,,". •
Chemical Senses. The chemical senses of taste
and smell are developed in varying degree. Accept-
ing the definition that what we perceive by the tongue
are tastes and the flavours we detect by the nose are
odours 112, insects can recognize the tastes of salt,
sweet, acid and bitter by means of receptors in the
mouth (Apis 97, &c.), on the antennae (Hymeno-
ptera) 189, 195, on the palpi (DytisGUS 225, Geotrupes 287,
&c.) and on the tarsi (Diptera, Lepidoptera) 183_
often in several of these sites in the same insect.
They can detect odours chiefly by sensilla on the
antennae 06, 1167, but to a lesser degree by the palps us.
While the' general chemical sense', affected by pun-
gent vapours, is present in many parts of the body 192.
Receptors for the perception of specific flavours occur
in the pharynx of sucking insects, notably the He-
miptera 268 ; thus the blood-sucking bug Rhodniuswill
rarely take up salt solution through a. membrane
unless a. trace of haemoglobin is added to it 268.
The sensitivity of the organs of taste in insects
may be very high: under conditions of starvation,
the butterfly Pyrameis, and the fly Oalliphora, may
SENSE ORGANS 109
be able to detect with their tarsi concentrations of
sugar of l~~bOO M or less 193; the threshold for the
human tongue lying between -h-h M. The apparent
sweetness of the various sugars may be very different
for insects and man 97; and insects may differ in
this respect not only between themselves, but even
when the organs of taste on the tarsi are compared
with those on the mouthparts: in Oalliphora, lactose
is apparently tasteless to the tarsi, but evokes a
response from the mouthparts 19&. Bya system of
rewards and penalties, the water beetle DytiaCU8 may
be trained to associate particular tastes with the
presence of food 226.
Insects vary enormously in the extent to which
their behaviour is directed by the sense of smell.
Many species find their food largely by this means
(cockroaches, crickets, Drosophila, &c. 289 A) ; butter-
flies often appear to be led to the food plant upon
which to lay their eggs by some single odorous con-
stituent in the leaves; parasitic Hymenoptera find
by smell the larva in which to la.y their eggs, some-
times detecting this from a distance, sometimes
following the trail of scent which it has left behind
it 199; the following of trails of scent is doubtless
very important also in the orientation of ants; but
the highest development of this sense is seen in the
males of those Lepidoptera (Lasiocampidae, &c.)
which can detect and locate the female from distances
of more than a mile by means of the olfactory organs
in the antennae 89.
The potentialities of the sense of smell have been
most fully studied in the case of the bee; and in
this respect the bee's perceptions seem to be surpris-
ingly like our own 96. Flowers odourless to us are
odourless to them; substances of unlike chemical
composition which smell alike to us-such as nitro-
benzol and oil of bitter almonds-are confused also
by the bee; and the threshold concentration for
odQurs to be detected seems to be of the same order
of magnitude for bee and man. Bees can readily
be trained to associate the presence of a supply of
food with particular scents; and they can retain the
memory of such associations for several weeks-far
longer than they can visual memories. But it is
interesting to note that they can be trained only
with aromatic odours such as might occur in flowers,
and not to such substances as indol or asafoetida 98.
Vision. We have already seen how varied are
the visual responses of insects. The receptor organs
which serve these responses range from small groups
of light. sensitive cells beneath lens-like thickenings
of the cuticle, to the compound eyes of the highest
forms, each with many thousands of retinal elements
and capable of tolerably good image perception.
The stemmata, or simple eyes of larvae, which
occur in the absence of compound eyes, are responsible
for the orientating responses to light in maggots 81
and caterpillars 170. In their highest development,
which is attained in the predaceous larvae of Cicin·
delids and other beetles, they serve to locate the
prey 92; and although it is doubtful if 'they are
capable of image perception, they can certainly per-
ceive the movement of objects within the field of
view. The ocelli, or simple eyes of adult insects, on
the other hand, are optically unsuited to eVen the
crudest type 6f image formation; for not only are
the retinal elements few in number, but any image
that is formed by the lens must fall far beyond the
level of what elements exist 294. In adult insects it
is the compound eyes which are responsible for the
movements to light: such movements do not occur
if the compound eyes are covered, and only the ocelli
remain 197.
What, then, is the function of the ocelli 1 So
far, the only suggestion of the many that have
been put forward 129, which has any experimental
evidence to support it, is that they are stimulatory
organs (p. 108) which accelerate phototaxis by in-
SENSE ORGANS 111
creasing the sensitivity of the brain to light-stimuli
received through the compound eyes. Thus the
response of Drosophila to light is appreciably more
rapid and more persistent if the ocelli are intact than
if they are blackened 31. It must be realized that
the light sense organs have two functions: a photo-
tactic or orientating function, such as we have already
discussed, and a photokinetic or stimulatory function
(well seen in many day-flying butterflies which become
quite incapable of flight in the absence of bright
sunlight 295). Now the phototactic response in
Drosophila is essentially an escape reaction, mani-
fested only when the insect is alarmed. The photo-
kinetic ocelli increase its efficiency.
But whenever a faceted compound eye is present,
no matter how few may be the retinal elements it
contains, these will inevitably receive different inten-
sities of light from different parts of the visual field;
and some. kind of visual pattern must always result.
Consequently, any movement in the visual field will
at once be perceived by the changes in pattern it wil1
produce. We have seen that many insects will turn
towards an object that is moving; will turn, that is,
until corresponding points of the two retinae are
stimulated by it. Now the closer the object is to
the insect, the nearer to the mid-line will be these
corresponding points. Consequently, the insect may
enjoy some of the advantages of binocular vision,
and be able to judge accurately the distance of objects,
even though its perception of images be rudimen-
tary 92; and like the larva of Aeschna 13 or the bug
Rhodniu8 289-' (when deprived of its antennae), it may
attack with precision any object that moves within
range.
In forms like the adult dragon-fly and the bee,
image perception reaches a higher plane, and specific
objects can be recognized; so that Sphingid moths,
for instance, will endeavour to extract nectar from
the flowers of wallpapers or from models of flowers 11H.
By calculation and by experiment it has been esti-
mated that the visual acuity of the bee, in the vertical
axis, is about one-fiftieth that of man (in the hori-
zontal axis the acuity is only about one-third of this,
because the eye is more curved and each ommatidium
has to cover a wider field-the bee, in fact, is astig-
matic 15, 118). But this degree of acuity is obtained
only under optimal conditions of illumination. With
poor illumination, both the visual acuity and the dis-
crimination of different light intensities 293 are so
greatly inferior to that of man, that the resolving
power of the eye must be extremely feeble. None
the less, these insects can recognize different forms,
and can learn to associate with these the presence
of food 95,297. The bee and other Hymenoptera can
find their way over miles of country by means of
landmarks 292, and can recognize the entrltnce to
their own hive or nest by visual memory 204, 255.
Colour Vision. The visual perceptions, notably
in flower-visiting insects, are greatly enhanced if the
objects are coloured; insects possess, in fact, a well-
developed colour vision 82,95. The histological basis
of this colour vision is not certainly known, though
it seems that each retinal element or ommatidium
receives a double innervation (comparable with the
rods and cones of vertebrates), one to perceive light
and the other to perceive colour 113. The functional
significance of colours in the vision of insects is much
better understood. Certain responses, such as the
fixation reflex, are influenced only by the luminOBity
of the source of stimulus and not by its colour 228.
Such responses, therefore, afford a method of testing
the sub~tive luminosity of different colours for
different insects; and it has been shown that for
many insects (Mantis, OocoineUa, Apis) the luminosity
of the visible spectrum is the same as for man with
the eye dark adapted; for others (pieNs) it corre-
sponds with the light adapted human eye 228. Many
insects can vary the amount of light entering the eye,
SENSE ORGANS 113
by movements of the pigment in the iris cells 206 ;
but whether this change has any influence on colour
vision is not known.
nor-------------------------------~
sso
500
¥SO
'1(10
l~JSO
.II!
~JOO
=I.!: 250
~
~
ZIJD
1.0
100
f6
6
706 ?SO
FIG. 13.-Relative stimulative efficiency of different wave

lengths for man and Drosophila. (After Bertholf.)
Corresponding with these differences in apparent

luminosity, there are differences in the visibility of
the spectrum. Certain insects (Apia 95; M'acroglo8sa,
Lep.) are practically blind to red, but can see far into
the ultra-violet 187. Such insects may be attracted
by the ultra-violet light reflected from certain
flowers 178; and they will cluster round screens
illuminated with ultra-violet rays which are invisible
to us 142. Indeed, the stimulating efficiency of the
ultra-violet part of the spectrum, for the honey-bee
and for Drosophila, may be far greater than that of
the regions visible to man (Fig. 13) 18, 224; and
although this effect may be due in some degree to
the fluorescence of parts of the eye under ultra-violet
illumination, this cannot account entirely for the
phenomenon 189. There must be a true perception
of ultra-violet; and it is interesting t9 note that the
limit of this perception is, approximately, the lower
limit of the solar spectrum 18. Other insects are
capable of seeing red (Pieridae) 140, and associated
with this is the shift in the apparent luminosity of
colours that we have already noted.
Insects, like man, experience the phenomenon of
simultaneous contrast; a grey field surrounded by
yellow appears blue to the bee, a grey field sur-
rounded by blue appears yellow 187.
Insect Behaviour. Most of the information on
colour vision in insects, and a great deal of the more
reliable information upon other senses, has been
obtained by training experiments: stimuli which the
insect can learn to associate with the presence of food,
or with the location of its nest, are judged to be per-
ceptible. Such experiments,. of course, need the
greatest care in their interpretation; for the insect
may be guided by some other stimulus that has been
overlooked by the experimenter; indeed, in the
course of a single experiment, it may cease to be
orientated by one stimulus and come to depend upon
another 292; and there are many who hold that the
whole perceptual experience of the organism is in-
tegrated, as it were, into a pattern, and that it
responds to general changes in this pattern, and not
BEHAVIOUR 115
to the isolated stimuli of which it is composed (Gestalt
theory) 93, 124.
Thus, when we consider the behaviour of insects
under natural conditions, we find them gaining their
ends by an infinite variety of sensory impressions.
Inborn reflexes, and reflexes 'conditioned' by the
experience of the individual, certainly occur; but
these are masked by inhibitions, and integrated by
higher centres, in such a way that they serve con-
stantly the needs and purposes of the insect as a
whole. It is this unifying quality, that welds the
organism together, and makes the whole something
greater than the sum of the parts, which in the sphere
of behaviour as in the sphere of growth still eludes
physiological analysis.
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116
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68 CLEVELAND, L. R. Quart. Rev. BioZ., 1 (1926), 51-60
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80 COUSIN, G. Bull. BioI. Fr. et Belg., Suppl., 15 (1932),
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88 Fox, H. M. Proo. Roy. Soc., B, 112 (1933), 479-95 (effect
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88 FBAENxEL, G. BioI. Roo., 6 (1931), 36--87 (mechanisms
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80 _ _ Z. tJ6rgl. Physiol., 16 (1932), 371-462 (nervous
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81 FREUDENSTEIN, K. Z. wiBs. Zooz., 132 (1928), 404-75
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81 FRIEDERIOHS, H. F. Z. Morph. Oekol. Piere, 21 (1931).
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88 _ _ Zool. Jahrb. Abe. Zool., 37 (1920), 1-238; 38 (1921),
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87 _ _ NaturwiBs., IS (1927), 321-7; 16 (1928), 307-15;
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88 FRIZA, F., and PRZIBB.AM, H. Arch. Entw. Mech., 53 (1933),
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88 GARREY, W. E. J. Gen. Physiol., 1 (1918). 101-25 (light
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100 GEnOULD, J. H. Quart. Rev. Biol., 3 (1927), 5S-78 (colours
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101 _ _ J. Morph., 48 (1929), 385-430; Science, 73 (1931,
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108 GIERBBERG, H. Z. vergl. Phllsio1., 7 (1928), 657-95 (colour
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108 _ _ Z. wrgl. Physiol., 9 (1929), 52~2 (determination
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108 _ _ BioI. BuZZ., 63 (1932), 337-56 (genetics and develop-
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107 GORTNER, R. A. Amer. Nat., 4S (1911), 743-55 (melanin
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108 GBED, R. J. BioI. BuZZ., 6S (1933), 179-86 (male mosaics
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111 HANSTROM, B. Z. vergE. Phyaiol., 4 (1926),528-44 (senses
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111 HAZELBOFF, E. H. Z. vergl. Phyaiol., 5 (1927), 179--90
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118 HELLER, J., et al. Biochem. Z., 219 (1930), 473-89; 255
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no HELLER, J., and AREM6WNA, H. Z. vergl. Phyaiol., 16
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123 HERTWECX, H. Z. wiBs. Zool., 139 (1931), 559--663
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114 HERTZ, M. Z. vergl. Physiol., 8 (1929), 693-748; 11
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125 HmsTAND, W. A. Physiol. Zool., 4 (1931), 246-70 (re-
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118 HnmER, A. Biol. Rev., 7 (1932), 224-53 (temperature
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117 HOBSON, R. P. J. Exp. Biol., 8 (1931), 109--23; Biochem.
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188 _ _ J. Exp. Biolo, 9 (1932), 12~38; 366-77; Biochem.
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181 HOLDAWA.Y, F. G., and SMITH, H. F. Auswal. J. Exp.
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181 HOLLANDE, A. C. Arch. Zool., 6 (1911), 283-323 (blood
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111__ Arch. Zool., 59 (1920), 543-63 (encapsulation of
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188 - _ Arch. d'Anat. Micr., 18 (1922), 85-307 (pericardial
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187 _ _ Arch. Zooz., 70 (1930), 231-80 (blood cells and
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139 Hoop, M. Zool. Jahrb. Abt. Anat., 57 (1933), 433-64
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144 KAESTNER. H. Arch. Entw. Mech., 124 (1931). 1-16 (in-
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148 KElLIN, D. Quart. J. Micr. Sci., 65 (1921), 611-25
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147 _ _ Ann. Mag. Nat. Hillt .• 13 (1924), 219-23 (pericardial
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148 KEMPER, H. Z. Morph. Oelcol. Tiere, 22 (1931), 53-109
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,148 - _ Z. Morph. Oelcot. Tiere. 15 (1929), 524-45 (stink
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160 KNIGHT, H. H. Ann. Ent. Soc. Amer., 17 (1924), 258-72
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161 KNOLL, F. Z. wrgl. PhysioZ .• 2 (1925), 329-80; Abhandl.
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187 _ _ Z. ver-gl. PhyBiol.• 5 (1927). 7~2-800 (colour vision:
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188 KUHNELT. W. Zool. Jah,.b. Abe. Anat.• 50 (1928). 219-
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189 KUNZE. G. Zool. Jah,.b. Abe. Zool .• 52 (1933). 465-512
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170 LAMMERT. A. Z. ve,.gl. Physiol.• 3 (19211). 225-78 (photic
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171 LANGE, H. H. Z. Zelljor-sch. u. mikr-. Anat.• 16 (1932),
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171 LAZARENKO, T. Z. mikr. Anat. Fo,.sch., 3 (1925), 409-99
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178 _ _ Z. mikr-. Anat. Forsch .• 12 (1928), 467-506 (healing
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174 v. LENGERXEN, H. Biol. Zent,.albl•• 44 (1924), 273-95
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178 LESTER, H. M. 0., and LLOYD, L. BuU. Ent. Rea., 19
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177 LOEB, J. Forced Movement8, Tropisms, and AnimaZ Con-
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178 LUDWIG, D. J. Ezp. Zooz., 60 (1931), 309-23 (respiratory
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180 McARTHUR, J. M. J. Ezp. Zool., 53 (1929), 117-28
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181l'11cGOVRAN, E. R. Ann. Ent. Soc. Amer., 24 (1931), 751-
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188 _ _ J. Ezp. Biol., 9 (1932), 222-31 (humidity and
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187 _ _ Proc. Ent. Soc. Lond., 1933, 22-4 (temperature of
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188 _ _ Proc. Roy. Soc., B (in press) (site of 'vater loss in
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188 MERKER, E. Zool. Jahrb. Abe. ZooZ., 46 (1929), 297-374;
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180 MEYER, E. Z. Morph. Oekol. Tiere, 22 (1931.), 1-52 (cir-
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181 MEYER. P. F. Z. vergZ. PhY8iol., 11 (1930), 173-209
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111 _ _ J. Ezp. Zool., 61 (1932), 375-93 (sense of taste in
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188MUXERJI, D. Ree. Ind. Mus., 34 (1932), 47-79 (excretory
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187 MULLEn, E. Z. wrgl. Physiol., 14 (1931),348-84 (function
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188 MiiLLER, G. W. Z. Morph. OekoZ. Tiere, 3 (1925). 542-
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100 Mur.rKOWSXI, R. A. Bull. Brooklyn Em. Soc., 19 (1924),
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101 NABERT, A. Z. wia8. ZooZ., 104 (1913), 181-358 (corpora
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131 SBAFElt, G. D. StanJord Uni". Publ. Biol. Sci .• 3 (1923),
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INDEX
Absorption, 55 Blood, osmotio pressure, 18,
Accessory hearts, 34, 35 37
Acenkopua,31 - pressure, 33
Acidia,63 - reo.ction of, 36
Aoiliua, 70 - specific gravity of, 36
Adipoleucocytes, 39 Blowfly, 51
AeBChna, 23, 24, 30, 33, 88, Bombyz, 36, 53
101, 110 Brain, 101
Agromyzidae, 62 Burmeister, 20
Air sacs, 22, 23, 30
- stores of aquatic insects, 28 Oalandra, 73
Akin\lBis, 103 Oalliphora, 53, 108
Aliform muscles, 33 Campanifoml sensilla, 108
Alimentary canal, 42 Oarabua,49
Amphipyra, 103 Carbon dioxide, elimination
Anaerobic metabolism, 77 of, 20
Anal gills, 30, 68 Cellulose digestion, 52
- papillae, 1,30,68 Cerambycidae, 62, 73
Anophele8, 48 Oeeonia,54
Anticoagulin, 49 Chemical senses, 108
Aorta, 32 Ohironomua, 29, 31
Apantele8, 31 Chitin, I, 8, 17
Api8, 33, 36, 37, 77, 108, - tests for, 2
109, 112, 113 Chordotonal organs, 107
ArtJBOhnia, 98 Chromogen, 12, 82
Athrooytes,57 Chrysomelidae, 14
Autotomy, 95 Ohrysops, 53
Oicindela, 49
Balkenlagen, 5 Cicindelida.e,4, 110
Behaviour, 100 Oimex, 45, 72, 88, 103
Binocular vision, III Circulation, mechanism of, 33
Blattidae, 46 Circulatory system, 32
Blood, cells, 38 Cirous movement, 102
- coagulation of, 37 Climbing organs, 'T
- composition of, 36 OloWn, 101
- functions of, 39 Clotting of blood, 37
- molecular concentration Coccids,14
of,37 Ooccin.ella, 112
130
INDEX 131
Cockroach, 13, 51, 71, 103, 109 Drosophilidae, 62
Cold resistance, 75 Dyar'slaw, 88
Collembola, 90 DytiBcus, 20, 22, 24, 28, 33,
ColoW' centre, 83, 99 35, 49, 103, 108
- change, morphological, 84,
95 Ecdysis, 7, 89
- - physiological, 83 Egg, hatching of, 13
- pattern, 95, 98 ElmiB,30
- vision, 112 Embiidae, 14
Ooretlira, 30, 101 Embioptera, 14
Oorixa, 26, 28, 29 Endocuticle, 1
Corpus allatum, 91 Energy metabolism, 77
Crew, 96 Enzymes, in gut, 61
Cricket, 109 - in salivary glands, 49
Crop, 44 Epicuticle, 2
Oryptooerous, 54 Eriogaster, 83
Cryptochaetum, 31, 63 EriBtaliB, 6, 20
Cuticle, composition of, 1 Evaporation from insects, 20,
- formation of, 9 74
- hardening and pigmenta- Excretion, 56
tion of, 11 - in aquatic insects. 68
- hydrophobe properties of, - of dyes, 56
6 - of nitrogen, 61
- lime in, 3, 60 - in Rhodnius, 63
- permeability of, 10 Exocuticle, 3
- stretching of, 5 Exuvial fluid, 8
- structure of. 4
Cuticulin, 2, 17 Fabre, 58
Fat body, 59, 63, 70
Deilephila, 36, 40 Fermentation chamber, 54
Dermal glands, 6, 8, 14 Filter chamber, 45
Desiccation, 74 Fixation reflex, 105
Diapause, 80, 9:1 Flea, 20, 46, 67
Diaphragms, 32 Food requirements, 7 J
Differentiation, 87 Forftcula, 48, 103
Diffusion control, 21
- in tracheae, 19 Galleria, 52, 87
Digestion, 42 Gastrophilus, 77
- extraintestinal, 50 Geotaxis, 107
Digestive enzymes, 51 Geotrup68, 108
Distance appreciation, III GerriB, 87
Di3:ippua, 20, 25, 82, 83, 87, Gestalt theory, 115
95 Gills, blood, 30
Donacia,31 - physical, 29
Dorow, 54 - rectal,30
Dormancy, 80 - tracheal, 30
Dragonfly, 13, 105 Glands, dermal, 6, 8, 14
Drosophila, 71, 87, 109, Ill, - labial, 49, 57
114 - moulting, 8, 60, 90
Glands, scent, 14 Image perception, III
- silk, 14, 69 Immunity, 40
- stink, 14 Intersexes, 96
- wax,14 Interventricular valves, 33
Glossina, 48, 53, 55, 72
Goldschmidt, 96, 98 Johnston's organ, 107
Graber, 22
Grabam,19 Kope~, 93
Granular leucocytes, :1Il Kowalevsky, 57
Growth, 80, 86 Krogh,19
- curves, 89
Gynundromorphs, 96 Lasiocampid80, 109
Gyrinidae, 106 LepiBma, 58, 65
Leptinotar8a, 74, 81
H abf'obracon, 83 Light reactions, 104
Haeckel,l Light-compass orientation,
Haematocytes,38 105
Haemoglobin, 41 Lime in cuticlll, 3, 60
Halteres, 108 - in Malpighian tubes, 60,
Hatching, 13 62,67
- spines, 13 Loeb,104
Healing of wounds, 38, 94 Lucilia, 52, 53, 72, 79
Hearing, 107 Lyrnantria, 93
Heart, 32 Lyonnet, 46
- nerve supply to, 35
Heart-beat, coritrol of, 35 MacroglosBa, 53, 113
- effect of carbon dioxide Malpighi, 16
on, 36 Malpighian tubes, 57, 60, 63
- revel'll8l of, 34 - - 8Ocessory functions, 68
- temperature coefficient of, Mansonia, 31
36 Mantia,1l2
Heteromorphous regenera- Mealworm, 73, 83
tion,95 Mechanical senses, 107
Hibernation, 74 Meconium, 61
Hind-gut, 65 Melanargia, 81
Histogenesis, 91, 93 Melanin, 11,82
Histolysis, 91, 93 Melaaorna, 15
Hopkins, 59 Melolontha, 22, 83
Hormones and growth, 81 Menotaxis, 105
- in metamorphosis, 92 Metabolism, 70
- in moulting, 90 Metamorphosis, blood cells
- and pigment movements, during, 38
84 - cause of, 91
- and sex determination, - chemical changes during,
97 79
Hydrophil'UB, 23, 28 Metapneustic respiration, 28,
Hydrophobe hairs, 7 31
Hydrostatic organs, 31 Milne Edwards, 58
Hypnosis, 103 Mosquito, 30, 31, 48, 68
INDEX 133
Moulting, 7, 89 Phototaxis, 111
- blood cells during, 38 Pieridae, 82
- cause of, 90 PuriB,112
- digestion of cuticle, 8 Pigments, 59, 81
- fluid, 8, 12 - of blood, 41
- - absorption of, 9 Plateau, 50
- glands,8 PopilUa, 53, 74
- mechanism of, 12 Pore-canals, 4, 12
Musca, 103 Poulton, 84
Mycetomes, 72 rroleucocytes, 38
Myrmeleon, 44 Prothetely, 93, 99
Proventriculus, 45
Nepa, 40 Przibram, 88
Nephrocytes, 57 Pseudonuclei,59
Nerve cord, 100 Ptilinum, 14
Nervous system, 100 Pulsatile organs, 35
Notonecta, 28, 88, 106 PulviIIi, 7
Nutrition, 70 Pupa, emergence from, 13
. - function of blood in, 39 Pupal metabolism, 78
Pupipara, 55, 72
Ocelli, 110 PyrameiB, 108
Oenocytes, 39, 70, 90
Oenocytoids, 39 Reaction of gut, 50
Oesophageal valve, 46 - of urine, 61
Orientation, 104 Rectal glands, 65
Oryctes, 54 Rectum, 65
Oamoderma, 54 Reflexes, 101, 102
Osmotic pressure of blood, Regeneration, 94
18,37 Reproduction, 86
Ostia, 32, 33 Respiration, 16
Oxygen supply, 19 - of aquatio insects, 28, 31
- cutaneous, 29, 31
Paedogenesis, 86 - function of blood in, 40
Parasitic castration, 97 - of parasites, 31
Parthenogenesis, 86 - regulation of, 24
Pediculus, 72 Respiratory centres, 21, 24,
Pericardial cells, 57 100
- sinus, 32 - metabolism, 76
Pericoma,3 - movements, 22
Perillus, 81 - quotient, 77
Perineural sinus, 32 Rhodnius, 4, 7, 8,45, 59, 63,
Periplaneta, 4, 25, 35, 53 88, 89, 92, 105, 108, 110
Peritrophic membrane, I, 46
Perivisceral sinus, .32, 35 Salivary glands, 49
Phagocytes, 38, 94 Samia,-83
Phagocytic organs, 38 Scent glands, 14
Phobotaxis, 103 SchiBtocerca, 24
Photinus, 105 Secondary sexual characterl!l,
Photokinesis, 11 1 96
Secretion, 50 Thigmotaxis, 103
Sekretschicht, 3 Tineola, 52, 89
Sense organs. 107 Topotaxis, 104
Sex detennination, 96 Touch, 107
Silk glands, 14, 69 Tracheae, 16
Silkwonn, 9, 51 - appearance of air in, 18
SimuUum, 29, 30 - penneability of, 17
Simultaneous contrast, 114 Tracheal capillaries, 17
Smell,108 - cell, 17
Sphinx, 36 - lungs, 40
Sphodromantis, 87 - system, ventilation of, 26
Spiracles, function of, 20, 24- Tracheoles, 16
Spiral filament, 15 - fluid in, 17
Stemmata, llO Treviranus, 19, 22, 106
Stereokinesis, 103, 107 Tropotaxis, 104
Stimulatory organs, 108, 110 Tsetse fly. 48, 49, 53, 55, 72
Stink glands, 14
Storage kidneys, 57 Ultra-violet perception, 114
Stratiomyiidae, 3, 62 Urate cells, 58
Stylop8,97 Uric acid, 58, 60, 61, 71
Suboesophageal ganglion, 101 Urine, 60
Supraoesophageal ganglion,
101 Van Wisselingh, 2
Surface forces and the cuticle, Ventilation control, 21
5 Verson's glands, 60, 90
Symbionts, in digestion, 54 Vesicular secretion, 50
- in nutrition, 72 Vespa, 35
Visceral nervous system, 35
Tabanu8,35 Visibility of spectrum, 113
Taenidia, 16 Vision, 105, 110
Taste. 108 Visual acuity, 112
Telotaxis, 105 - intensity discrimination,
Temperature control, 74 112
- of insects, 75 , Vitamins, 72
- resistance, 74
Tenebrio, 4, 66, 79, 83, 89, Water metabolism, 73
93,94 Wax glands, 14
Tension receptors, 107 WeisnnaIU1, 38, 57
'rennites, 54 Wings, expansion of, 13,33
Thanatosis, 103 Wound healing. 38. 94-
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Insect Physiology (Wigglesworth)

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INSECT PHYSIOLOGY

METHUEN & CO. LTD.

T HE fundamental ·processes of vital activity,

III THE CIRCULATORY SYSTEM .AND THE BLOOD 32

VI NUTRITION AND METABOLISM. 70

VII REPRODUCTION AND GROWTH • 86

VIII THE NERVOUS SYSTEM, SENSE ORGANS AND

T HE key to much of the physiology of insects is

layer is responsible for the rigidity of the cuticle ;

FIG. I.-The insect cuticle: semischematic

times it is laid down, before moulting, in pits in the

Fig. 2.-Diagrams illustrating the properties of the inseot

produoing a fatty secretion 2'. That is the case, for

meability when mature. For we have seen that the

dOWnwards towards the tissues (Fig. 4, B). And if

N the insect body there is only one tissue fluid,

simultaneously and extinguish one another when

perhaps because these take only liquid food, and in

T HE function of the excretory organs of animals

FIG. 9.-A, excretory system of RhodniuB: schematic;

appears above the upper ligature and below * the

FIG. lO.-Types of excretory system in insects: schematic

dry substances 210, such as the mealworm (Tenehrio),

INCE the alimentary canal of insects contains

metabolism is a measure of the amount of formed

T HE reproduction of insects forms one of the

increase in size (p. 12). ThE)re is no doubt, therefore,

FIG. 13.-Relative stimulative efficiency of different wave

Corresponding with these differences in apparent

THE MECHANISM AND PHYSIOLOGY OF SEX DETERMINA-

THE OPPOSITE SEXES r A Bloloatcal and Psycholoatcal Study

SCIENCE FROM AN EASY CHAIR: First Series.

METHUEN'S MONOGRAPHS ON BIOLOGICAL

THE SNAKES OF EUROPE

A GENERAL TEXT-BOOK OF ENTOMOLOGY

A TEXT-BOOK OF PLANT BIOLOGY

HOW TO KNOW THE FERNS

PLANT PARASITIC NEMATODES, aad the Diseases they cause

METHUEN'S GEOLOGICAL SERIES

THE NAPPE THEORY IN THE ALPS: Alpine Tectonfc:s. 1905-1928

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