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 Another Random Document on
Scribd Without Any Related Topics
 The Alimentary Canal.
3
The alimentary canal of the Cockroach measures about 2 4 inches in
3
length, and is therefore about 2 4 times the length of the body. In
herbivorous Insects the relative length of the alimentary canal may
be much greater than this; it is five times the length of the body in
Hydrophilus. Parts of the canal are specialised for different digestive
offices, and their order and relative size are given in the following
table:—
Œsophagus and crop ·95 in.
Gizzard ·1
Chylific stomach ·5
Small intestine ·1
Colon ·875
Rectum ·25
────
2·775
════

The principal appendages of the alimentary canal are the salivary

glands, the cæcal diverticula of the stomach, and the Malpighian
tubules.
Considered with respect to its mode of formation, the alimentary
canal of all but the very simplest animals falls into three sections—
viz., (1) the mesenteron, or primitive digestive cavity, lined by
hypoblast; (2) the stomodæum, or mouth-section, lined by epiblast,
continuous with that of the external surface; and (3) the
proctodæum, or anal section, lined by epiblast folded inwards from
the anus, just as the epiblast of the stomodæum is folded in from
 the mouth. The mesenteron of the
Cockroach is very short, as in other
Arthropoda, and includes only the
chylific stomach with its diverticula.
The mouth, œsophagus, and crop
form the stomodæum, while the
proctodæum begins with the
Malpighian tubules, and extends
thence to the anus. Both
stomodæum and proctodæum have
a chitinous lining, which is wanting
in the mesenteron. At the time of
moult, or a little after, this lining is
broken up and passed out of the
body.
The mouth of the Cockroach is
enclosed between the labrum in
front, and the labium behind, while
it is bounded laterally by the
Fig. 56.—Alimentary Canal mandibles and first pair of maxillæ.
of Cock­roach. × 2. The chitinous lining is thrown into
many folds, some of which can be
obliterated by distension, while others are permanent and filled with
solid tissues. The lingua is such a permanent fold, lying like a tongue
upon the posterior wall of the cavity and reaching as far as the
external opening. The thin chitinous surface of the lingua is hairy,
like other parts of the mouth, and stiffened by special chitinous rods
or bands. The salivary ducts open by a common orifice on its hinder
surface. Above, the mouth leads into a narrow gullet or œsophagus,
with longitudinally folded walls, which traverses the nervous ring,
and then passes through the occipital foramen to the neck and
thorax. Here it gradually dilates into the long and capacious crop,
whose large rounded end occupies the fore-part of the abdomen.
When empty, or half-empty, the wall of the crop contracts, and is
thrown into longitudinal folds, which disappear on distension.
Numerous tracheal tubes ramify upon its outer surface, and appear
as fine white threads upon a greenish-grey ground.

Fig. 57.—Section of Wall of Crop.

Cc, chi­tin­ous layer; C, chi­tin­‐
ogenous cells; Mi, inner mus­cular Fig. 58.—Wall of
layer; Mo, outer do. × 275. Crop, in suc­ces­sive
lay­ers. Refer­ences
as in fig. 57.
× 250.

Three layers can be distinguished in the wall of the crop—viz., (1)

the muscular, (2) the epithelial, and (3) the chitinous layer. 122 The
muscular layer consists of annular and longitudinal fibres, crossing at
right angles. (See fig. 58.) In most animals the muscles of organic
life, subservient to nutrition and reproduction, are very largely
composed of plain or unstriped fibres. In Arthropoda (with the
exception of the anomalous Peripatus) this is not generally the case,
and the muscular fibres of the alimentary canal belong to the striped
variety. The epithelium rests upon a thin structureless basement-
membrane, which is firmly united in the œsophagus and crop to the
muscular layer and the epithelium. The epithelium consists of
scattered nucleated cells, rounded or oval. These epithelial cells,
homologues of the chitinogenous cells of the integument, secrete
the transparent and structureless chitinous lining. Hairs (setæ) of
elongate, conical form, and often articulated at the base, like the
large setæ of the outer skin, are abundant. In the œsophagus they
are very long, and grouped in bundles along sinuous transverse
lines. In the crop the hairs become shorter, and the sinuous lines run
into a polygonal network. The points of the hairs are directed
backwards, and they no doubt serve to guide the flow of saliva
towards the crop.
 Fig. 59.—Transverse section
of Giz­zard of Cock­roach. The
chi­tin­ous folds are rep­re­sent­‐
ed here as sym­metri­cal. See
next figure. × 30.

Fig. 60.—The Six Primary Folds (teeth) of

the Giz­zard, seen in pro­file.

The gizzard has externally the form of a blunt cone, attached by its
base to the hinder end of the crop, and produced at the other end
1 1
into a narrow tube ( 4 to 3 in. long), which projects into the chylific
stomach. Its muscular wall is thick, and consists of many layers of
annular fibres, while the internal cavity is nearly closed by radiating
folds of the chitinous lining. Six of the principal folds, the so-called
“teeth,” are much stronger than the rest, and project so far inwards
that they nearly meet. They vary in form, but are generally
triangular in cross section and irregularly quadrilateral in side view.
Between each pair are three much less prominent folds, and
between these again are slight risings of the chitinous lining. A ridge
runs along each side of the base of each principal tooth, and the
minor folds, as well as part of the principal teeth, are covered with
fine hairs. The central one of each set of secondary folds is produced
behind into a spoon-shaped process, which extends considerably
beyond the rest, and gradually subsides till it hardly projects from
the internal surface of the gizzard. Behind each large tooth (i.e.,
towards the chylific stomach) is a rounded cushion set closely with
hairs, and between and beyond these are hairy ridges. (See fig. 61.)
The whole forms an elaborate machine for squeezing and straining
the food, and recalls the gastric mill and pyloric strainer of the
Crayfish. The powerful annular muscles approximate the teeth and
folds, closing the passage, while small longitudinal muscles, which
can be traced from the chitinous teeth to the cushions, appear to
retract these last, and open a passage for the food. 123
Fig. 61.—Part of Gizzard laid open,
show­ing two teeth (T) and the
inter­medi­ate folds, as well as the
hairy pads below. A-A and B-B are
lines of sec­tion (see figs. 62 and
63). × 50.
 Fig. 62.—Section through one tooth and
two inter­medi­ate spa­ces (see figure 61, A-
A). Cc, chi­tin­ous cuti­cle; C, chi­tin­ogenous
layer; am, annu­lar mus­cles; p, peri­ton­eal
layer. × 75.

Fig. 63.—Section through one

princi­pal hairy ridge and two inter­‐
medi­ate spaces (see fig. 61, B-B);
rm, radi­at­ing mus­cles; tr, tra­chea.
The other refer­ences as before.
× 75.

The gizzard ends below, as we have already mentioned, in a narrow

cylindrical tube which is protruded into the chylific stomach for about
one-third of an inch. Folds project from the wall of this tube, and
reduce its central cavity to an irregular star-like figure. Below it ends
in free processes slightly different from each other in size and shape.
The chitinous lining and the chitinogenous layer beneath pass to the
end of the tube and are then reflected upon its outer wall, ascending
till they meet the lining epithelium of the cæcal tubes. Between the
wall of the gizzard-tube and its external reflected layer, tracheal
tubes, fat-cells, and longitudinal muscles are enclosed.
Fig. 64.—Longitudinal section through Giz­zard and
fore-part of Chyl­ific Stom­ach. G, giz­zard; Tu, cæc­al
tube; St, stom­ach; Ep, its lin­ing epi­the­lium. A and B
are en­larged in the side fig­ures. × 35.
A.—The Reflected Chitinogenous Layer of the
Tubular Gizzard. Tr, tracheal tube. × 400.
B.—One of the Tubular Extensions of the same,
enclosing muscles and tracheæ. × 400.
The chylific stomach is a simple cylindrical tube, provided at its
anterior end with eight (sometimes fewer) cæcal tubes, and opening
behind into the intestine. Its muscular coat consists of a loose layer
of longitudinal fibres, enclosing annular fibres. Internal to these is a
basement membrane, which supports an epithelium consisting of
elongate cells which are often clustered into regular eminences, and
separated by deep cavities. The epithelium forms no chitinous lining
in the chylific stomach or cæcal tubes; and this peculiarity, no doubt,
promotes absorption of soluble food in this part of the alimentary
canal. Short processes are given off from the free ends of the
epithelial cells, as in the intestines of many Mammalia and other
animals.
Fig. 65.—Transverse section of tubu­‐
lar pro­long­ation of Giz­zard, with­in
the Chyl­ific Stom­ach, part of which
is shown at its pro­per dis­tance. R C,
re­flect­ed chi­tin­ogenous layer; Tr,
tra­cheal tube; M, cross sec­tion of
mus­cle; Ep, epi­the­lium of chyl­ific
stom­ach. × 100.
 Fig. 66.—Epithelium of
Chylific Stom­­ach. In the
upper figure the di­gest­ive
sur­face is in­dent­ed, while in
the lower fig­ure it is flat.
Both ar­range­ments are com­‐
mon, and may be seen in a
single sec­tion. The epi­the­lial
buds are shown below, and
again below these the annu­‐
lar and longi­tud­inal mus­cles.
× 220.

Between the cells a reticulum is often to be seen, especially where

the cells have burst; it extends between and among all the elements
of the mucous lining, and probably serves, like the very similar
structure met with in Mammalian intestines, 124 to absorb and
conduct some of the products of digestion. Different epithelial cells
may be found in all the stages noticed by Watney—viz., (1) with
divided nuclei; (2) small, newly produced cells at the base of the
epithelium; (3) short and broad cells, overtopped by the older cells
around; (4) dome-shaped masses of young cells, forming “epithelial
buds”; 125 (5) full-grown cells, ranging with those on either side, so
as to form an unbroken and uniform series. The regeneration of the
tissue is thus provided for. The cells come to maturity and burst,
when new cells, the product of the epithelial buds, take their place.
The epithelium of the chylific stomach is continued into the eight
cæcal tubes, where it undergoes a slight modification of form.
At the hinder end of the chylific stomach is a very short tube about
half the diameter of the stomach, the small intestine. At its junction
with the chylific stomach are attached, in six bundles, 60 or 70 long
and fine tubules, the Malpighian tubules. 126 The small intestine has
the same general structure as the œsophagus and crop; its chitinous
lining is hairy, and thrown into longitudinal folds which become much
more prominent in the lower part of the tube. The junction of the
small intestine with the colon is abrupt, and a strong annular fold
assumes the character of a circular valve (fig. 68).
Fig. 67.—Section of Chylific Stomach, show­ing the
six bund­les of Mal­pigh­ian tub­ules. × 70.
 Fig. 68.—Junction of
Small In­test­ine with
Colon. × 15.

From the circular valve the colon extends for nearly an inch. Its
diameter is somewhat greater than that of the chylific stomach, and
uniform throughout, except for a lateral diverticulum or cæcum,
which is occasionally but not constantly present towards its rectal
end. The fore part of the colon is thrown into a loose spiral coil. A
constriction divides the colon from the next division of the
alimentary canal, the rectum.
1
The rectum is about 4
inch long, and is dilated in the middle when
distended. Six conspicuous longitudinal folds project into the lumen
of the tube. These folds are characterised by an unusual
development of the epithelium, which is altogether wanting in the
intermediate spaces, where the chitinous lining blends with the
basement-membrane, both being thrown into sharp longitudinal
corrugations. Between the six epithelial bands and the muscular
layer are as many triangular spaces, in which ramify tracheal tubes
and fine nerves for the supply of the epithelium. The chitinous layer
is finely setose. The muscular layer consists of annular fibres
strengthened externally by longitudinal fibres along the interspaces
between the six primary folds. 127
Fig. 69.—Transverse section of Small
Intestine and Colon, close to their junction.
× 50.

Fig. 70.—Transverse section of

Rectum. × 50.
The corrugated and non-epitheliated interspaces may be supposed
to favour distension of the rectal chamber, while the great size of the
cells of the bands of epithelium is perhaps due to their limited
extent. Leydig 128 attributed to these rectal bands a respiratory
function, and compared them to the epithelial folds of the rectum of
Libellulid larvæ, which, as is well known, respire by admitting fresh
supplies of water into this cavity. It is an obvious objection that
Cockroaches and other Insects in which the rectal bands are well
developed do not take water into the intestine at all. Gegenbaur has
therefore modified Leydig’s hypothesis. He suggests (Grundzüge d.
Vergl. Anat.) that the functional rectal folds of Dragon-flies and the
non-functional folds of terrestrial Insects are both survivals of
tracheal gills, which were the only primitive organs of respiration of
Insects. The late appearance of the rectal folds and the much earlier
appearance of spiracles is a serious difficulty in the way of this view,
as Chun has pointed out. It seems more probable that the
respiratory appendages of the rectum of the Dragon-fly larvæ are
special adaptations to aquatic conditions of a structure which
originated in terrestrial Insects, and had primarily nothing to do with
respiration.
The number of the rectal bands (six) is worthy of remark. We find
six sets of folds in the gizzard and small intestine of the Cockroach,
six bundles of Malpighian tubules, with six intermediate epitheliated
bands. There are also six longitudinal bands in the intestine of the
Lobster and Crayfish. The tendency to produce a six-banded
stomodæum and proctodæum may possibly be related to the six
theoretical elements (two tergal, two pleural, two sternal,) traceable
in the Arthropod exoskeleton, of which the proctodæum and
stomodæum are reflected folds.
The anus of the Cockroach opens beneath the tenth tergum, and
between two “podical” plates. Anal glands, such as occur in some
Beetles, have not been discovered in Cockroaches.
 Appendages. The Salivary Glands.
The three principal appendages of the alimentary canal of the
Cockroach are outgrowths of the three primary divisions of the
digestive tube; the salivary glands are diverticula of the
stomodæum, the cæcal tubes of the mesenteron, and the
Malpighian tubules of the proctodæum.

Fig. 71.—Salivary Glands and Receptacle, right side. The

arrow marks the open­ing of the com­mon duct on the back
of the lingua. A, side view of lingua; B, front view of
lingua.

A large salivary gland and reservoir lie on each side of the

1
œsophagus and crop. The gland is a thin foliaceous mass about 3 in.
long, and composed of numerous acini, which are grouped into two
principal lobes. The efferent ducts form a trunk, which receives a
branch from a small accessory lobe, and then unites with its fellow.
The common glandular duct thus formed opens into the much larger
common receptacular duct, formed by the union of paired outlets
from the salivary reservoirs. The common salivary duct opens
beneath the lingua. Each salivary reservoir is an oval sac with
transparent walls, and about half as long again as the gland. The
ducts and reservoirs have a chitinous lining, and the ducts exhibit a
transverse marking like that of a tracheal tube. When examined with
high powers the wall of the salivary gland shows a network of
protoplasm with large scattered nuclei, resting upon a structureless
chitinous membrane.
The salivary glands are unusually large in most Orthoptera. 129 In
other orders they are of variable occurrence and of very unequal
development.

The Cæcal Tubes.

There are eight (sometimes fewer) cæcal tubes arranged in a ring
round the fore end of the chylific stomach; they vary in length, the
longer ones, which are about equal to the length of the stomach
itself, usually alternating with shorter ones, though irregularities of
arrangement are common. The tubes are diverticula of the stomach
and lined by a similar epithelium. In the living animal they are
sometimes filled with a whitish granular fluid.
Similar cæcal tubes, sometimes very numerous and densely
clustered, are attached to the stomach in many Crustacea and
Arachnida. The researches of Hoppe Seyler, Krukenberg, Plateau,
and others have established the digestive properties of the fluid
secreted in them, which agrees with the pancreatic juice of
Vertebrates.
 The Malpighian Tubules.
The Malpighian tubules mark the beginning of the small intestine, to
which they properly belong. They are very numerous (60–70) in the
Cockroach, as in Locusts, Earwigs, and Dragon-flies; and
unbranched, as in most Insects. They are about ·8 inch in length,
and ·002 inch in transverse diameter, so that they are barely visible
to the naked eye as single threads. In larvæ about one-fifth of an
inch long, Schindler 130 found only eight long tubules, the usual
number in Thysanura, Anoplura, and Termes; but the grouping into
six masses, so plainly seen in the adult, throws some doubt upon
this observation. In the adult Cockroach the long threads wind about
the abdominal cavity and its contained viscera.
In the wall of a Malpighian tubule
there may be distinguished (1) a
connective tissue layer, with fine
fibres and nuclei; within this, (2) a
basement-membrane, between
which and the connective tissue layer
runs a delicate, unbranched tracheal
tube; (3) an epithelium of relatively
large, nucleated cells, in a single Fig. 72.—Malpighian
layer, nearly filling the tube, and Tubules of Cock­roach. A,
leaving only a narrow, irregular trans­verse sec­tion of
central canal. Transverse sections young tubule; p, its con­‐
show from four to ten of these cells nect­ive-tissue or “peri­ton­‐
at once. The tubules appear eal” layer; B, older tubule,
transparent or yellow-white, crowd­ed with urates; tr,
according as they are empty or full; tra­cheal tube; C, tubule
sometimes they are beaded or cut open longi­tudi­nal­ly,
varicose; in other cases, one half is show­ing three states of
coloured and the other clear. The
opaque contents consist partly of the lin­ing epi­the­lium.
crystals, which usually occur singly in × 200.
the epithelial cells, or heaped up in
the central canal. Occasionally, they form spherical concretions with
a radiate arrangement. They contain uric acid, and probably consist
of urate of soda. 131 In the living Insect the tubules remove urates
from the blood which bathes the viscera; the salts are condensed
and crystallised in the epithelial cells, by whose dehiscence they
pass into the central canals of the tubules, and thence into the
intestine.
The Malpighian tubules develop as diverticula from the proctodæum,
which is an invagination of the outer integument and its
morphological equivalent. They are, therefore, similar in origin to
urinary organs opening upon the surface of the body and developed
as invaginations of the integument, like the “shell-glands” of lower
Crustacea, and the “green glands” of Decapod Crustacea. The
segmental organs of Peripatus, Annelids, and Vertebrates do not
appear to be possible equivalents of the excretory organs of
Arthropods. They arise, not as involutions, but as solid masses of
mesoblastic tissue, or as channels constricted off from the peritoneal
cavity, and their ducts have only a secondary connection with the
outside of the body or with the alimentary canal.

Digestion of Insects.
The investigation of the digestive processes in Insects is work of
extreme difficulty, and it is not surprising that much yet remains to
be discovered. Plateau has, however, succeeded in solving some of
the more important questions, which, before his time, had been
dealt with in an incomplete or otherwise unsatisfactory way. The
experiments of Basch, though now superseded by Plateau’s more
trustworthy results, deserve notice as first attempts to investigate
the properties of the digestive fluids of Insects.
Basch set out with a conviction that where a chitinous lining is
present, the epithelium of the alimentary canal secretes chitin only,
and that proper digestive juices are only elaborated in the chylific
stomach, or in the salivary glands. The tests applied by him seemed
to show that the saliva, as well as the contents of the œsophagus
and crop, had an acid reaction, while the contents of the chylific
stomach were neutral at the beginning of the tube and alkaline
further down. From this he concluded that the supposed deep-
seated glands of the chylific stomach secreted an alkaline fluid,
which neutralised the acidity of the saliva. Finding that the epithelial
cells of the stomach were often loaded with oil-drops, he concluded
that absorption, at least of fats, takes place here. The chylific
stomach, carefully emptied of its contents, was found to convert
starch into sugar at ordinary temperatures. The saliva of the
Cockroach gave a similar result, and when a weak solution of
hydrochloric acid was added, Basch thought that the mixture could
digest blood-fibrin at ordinary temperatures.
Plateau’s researches upon Periplaneta americana, 132 modified by
subsequent experiments upon P. orientalis, 133 and by still more
recent observations, lead him to the following conclusions 134:—
1.—The saliva of the Cockroach changes starch into glucose; but the
saliva is not acid, it is either neutral (P. orientalis) or alkaline (P.
americana). Any decided acidity found in the crop is due to the
ingestion of acid food; but a very faint acidity may occur, which
results from the presence in the crop of a fluid secreted by the cæcal
diverticula of the mesenteron.
2.—The glucose thus formed is absorbed in the crop, and no more is
formed in the succeeding parts of the digestive tube.
3.—The function of the gizzard is that of a grating or strainer. It has
no power of trituration. If the animal consumes vegetable food rich
in cellulose, a substance not capable of digestion in the crop, the
fragments are found unaltered as to form and size in the
mesenteron. If it is supplied with plenty of farinaceous food, such as
meal or flour, the saliva is not adequate to the complete solution and
transformation of the starch, and the intestine is found full of
uninjured starch granules, which must have traversed the gizzard
without crushing.
4.—The cæcal diverticula secrete a feebly acid fluid. To demonstrate
its acidity an extremely sensitive litmus solution, capable of
indicating one part in twenty thousand of hydrochloric acid, must be
used. The fluid secreted by the cæca emulsifies fats, and converts
albuminoids into peptones.
In all Insects digestion is effected in the following way (which is
particularly easy of demonstration in Carabus and Dytiscus). The
crop is filled with food coarsely divided by the mandibles, and the
gizzard being shut to prevent further passage, the fluid secretion of
the cæca ascends to the crop, and there acts upon the food.
Digestion is effected in the crop, and not beyond it. This is clear
beyond doubt. In Decapod Crustacea also it is very easy to prove
that the fluid secreted by the so-called liver ascends into the
stomach (which corresponds to the crop, together with the gizzard
of the Insect). To satisfy ourselves on this point we have only to
open a Crayfish during active digestion.
When digestion in the crop is finished, the gizzard relaxes, and the
contents of the crop, now in a semi-fluid condition, pass into the
mesenteron, which is devoid of chitinous lining, and particularly
fitted for absorption.
5.—There are no absorbent vessels properly so called, and Plateau
has long thought that the products of digestion pass by osmosis
directly through the walls of the digestive tube, to mix with the
blood in the perivisceral space. If we may rely upon what is now
known of the process in Vertebrates, we should be led to modify this
explanation. It is very likely that in Insects, as in Vertebrates,
absorption is effected by the protoplasm of the epithelial cells, which
select and appropriate certain substances formed out of the
dissolved food. Not only do the epithelial cells transmit to the
neighbouring blood-currents the materials which they have
previously absorbed, but they subject certain kinds to further
elaboration. The protoplasm of the epithelial cells of Vertebrates is
capable of forming fat. Thus, a mixture of soap and glycerine,
injected into the intestine of a Vertebrate, is absorbed by the lacteals
in the form of oil-drops. Modern physiologists allow, too, that part of
the peptone is similarly changed into albumen, without transport to
a distance, by the activity of the epithelial lining.
These facts explain why Plateau was unable to isolate the secretion
of the epithelium of the chylific stomach of Insects. The cells are not
secretory, but absorbent; and the secretion vainly sought for does
not actually exist.
 CHAPTER VIII.
The Organs of Circulation and Respiration.
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