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Discovering Diverse Content Through
Random Scribd Documents
 SPECIAL REFERENCES.
Verloren. Mém. sur la Circulation dans les Insectes. Mém. cour, par l’Acad.
Roy. de Belgique, Tom. XIX. (1847). [Structure of Circulatory Organs in a
number of different Insects.]
Graber. Ueb. den Propulsatorischen Apparat der Insekten. Arch. f. mikr.
Anat., Bd. IX. (1872). [Heart and Pericardium.]
Leydig. Larve von Corethra plumicornis. Zeits. f. wiss. Zool., Bd. III.
(1852). [Valves in Heart.]
Landois, H. Beob. üb. das Blut der Insekten. Zeits. f. wiss. Zool., Bd. XIV.
(1864). [Blood of Insects.]
Jaworowski. Entw. des Rückengefässes, &c., bei Chironomus. Sitzb. der k.
Akad. der Wiss. Wien., Bd. LXXX. (1879). [Minute Structure and
Development of Heart.]
Landois, H., and Thelen. Der Tracheenverschluss bei den Insekten. Zeits. f.
wiss. Zool., Bd. XVII. (1867). [Stigmata.]
Palmen. Zur Morphologie des Tracheensystems (1877). [Morphology of
Stigmata and Tracheal Gills.]
MacLeod. La Structure des Trachées et la Circulation Péritrachéenne.
(Brussels, 1880.)
Lubbock. Distribution of Tracheæ in Insects. Trans. Linn. Soc., Vol. XXIII.
(1860).
Rathke. Untersuch. üb. den Athmungsprozess der Insekten. Schr. d. Phys.
Oek. Gesellsch. zu Königsberg. Jahrg. I. (1861). [Experiments and
Observations on Insect-respiration.]
Plateau. Rech. Expérimentales sur les Mouvements Respiratoires des
Insectes. Mém. de l’Acad. Roy. de Belgique, Tom. XLV. (1884). Preliminary
notice in Bull. Acad. Roy. de Belgique, 1882.
Langendorff. Studien üb. die Innervation der Athembewegungen.—Das
Athmungscentrum der Insekten. Arch. f. Anat. u. Phys. (1883).
[Respiratory Centres of Insects.]

Circulation of Insects.
A very long chapter might be written upon the views advanced by
different writers as to the circulation of Insects. Malpighi first
discovered the heart or dorsal vessel in the young Silkworm. His
account is tolerably full and remarkably free from mistakes. The
heart of the Silkworm, he tells us, extends the whole length of the
body, and its pulsations are externally visible in young larvæ. He
supposed that contraction is effected by muscular fibres, but these
he could not distinctly see. The tube, he says, has no single large
chamber, but is formed of many little hearts (corcula) leading one
into another. The number of these he could not certainly make out,
but believed that there was one to each segment of the body. During
contraction each chamber became more rounded, and when
contraction was specially energetic, the sides of the tube appeared
to meet at the constrictions. The flow of blood, he ascertained, was
forward, the rhythm not constant. No arteries were seen to be given
off from the heart. 135 Swammerdam thought that his injections
ascertained the existence of vessels branching out from the heart, 136
but this proved to be a mistake. Lyonnet added many details of
interest to what was previously known. He came to the conclusion
that there was no system of vessels connected with the heart, and
even doubted whether the organ so named was in effect a heart at
all. Marcel de Serres maintained that it was merely the secreting
organ of the fat-body. Cuvier and Dufour doubted whether any
circulation, except of air, existed in Insects. This was the extreme
point of scepticism, and naturalists were drawn back from it by
Herold, 137 who repeated and confirmed the views held by the
seventeenth-century anatomists, and insisted upon the
demonstrable fact that the dorsal vessel of an Insect does actually
pulsate and impel a current of fluid. Carus, in 1826, saw the blood
flowing in definite channels in the wings, antennæ, and legs. Straus-
Durckheim followed up this discovery by demonstrating the
contractile and valvular structures of the dorsal vessel. Blanchard
affirmed that a complex system of vessels accompanied the air tubes
throughout the body, occupying peritracheal spaces supposed to
exist between the inner and outer walls of the tracheæ. This
peritracheal circulation has not withstood critical inquiry, 138 and it
might be pronounced wholly imaginary, except for the fact that air
tubes and nerves are found here and there within the veins of the
wings of Insects.
Fig. 73.—Heart, Alary Muscles, and
Tracheal Arches, seen from below; to the
left is a side view of the heart. T2, T3, A1,
alary mus­cles attached to the sec­ond
 thora­cic, third thora­cic, and first ab­dom­‐
inal terga. × 6. Fig. 35 (p. 74) is not
quite cor­rect as to the details of the
heart. The thor­acic por­tion should be
cham­bered, and add­itional cham­bers and
alary muscles rep­re­sent­ed at the end of
the ab­do­men. These omis­sions are recti­‐
fied in the pres­ent fig­ure.

Heart of the Cockroach.

The heart of the Cockroach is a long, narrow tube, lying immediately
beneath the middle line of the thorax and abdomen. It consists of
thirteen segments (fig. 73), which correspond to three thoracic and
ten abdominal somites. Each segment, as a rule, ends behind in a
conspicuous fold which projects backwards from the dorsal surface;
immediately in front of this are two lateral lobes. The median lobe
passes into the angle between two adjacent terga, and is continuous
with the dorsal wall of the segment next behind, from which it is
separated only by a deep constriction, while the lateral folds conceal
paired lateral inlets, 139 which lead from the pericardial space to the
hinder end of each chamber of the heart. Immediately in front of
each constriction is the interventricular valve, a pear-shaped mass of
nucleated cells, hanging down from the upper wall of the heart, and
inclining forward below. The position of this valve indicates that
during systole it closes upon the constricted boundary between two
chambers, thus shutting off at once the inlets and the passage into
the chambers behind. In this way the progressive and rhythmical
contraction of the chambers impels a steady forward current of
blood, allowing an intermittent stream to enter from the pericardial
space, but preventing regurgitation.
 Fig. 74.—Diagram to
show the inter­ven­tric­ular Fig. 75.—
valves and lat­eral inlets of Junction of two
the Heart. ML, med­ian chambers of the
lobe; V, valve; I, lat­eral Heart, seen
inlet. from above. ML,
median lobe; I,
lateral inlet.

The wall of the heart includes several distinct layers. There are (1) a
transparent, structureless intima, only visible when thrown into
folds; (2) a partial endocardium, of scattered, nucleated cells, which
passes into the interventricular valves; (3) a muscular layer,
consisting of close-set annular, and distant longitudinal fibres. The
annular muscles are slightly interrupted at regular and frequent
intervals, and are imperfectly joined along the middle line above and
below, so as to indicate (what has been independently proved) that
the heart arises as two half-tubes, which afterwards join along the
middle. Elongate nuclei are to be seen here and there among the
muscles. The adventitia (4), or connective tissue layer, is but slightly
developed in the adult Cockroach.
Within the muscular layer is a structure which we have failed to
make out to our own satisfaction. It presents the appearance of
regular but imperfect rings, which do not extend over the upper
third of the heart. They probably meet in a ventral suture, but this
and other details are hard to make out, owing to the transparency of
the parts. The rings stain with difficulty, and we have not observed
nuclei belonging to them. Each extends over more than one bundle
of annular muscles.
The difficulty of investigating a structure so minute and delicate as
the heart of an Insect may explain a good deal of the discrepancy
noted on comparing various published descriptions. Perhaps the
most obvious peculiarity which distinguishes the heart of the
Cockroach, is the subdivision of the thoracic portions into three
chambers, which, though less prominent in side-view than the
abdominal chambers, are, nevertheless, perfectly distinct. The
number of abdominal chambers is also unusually high; but it is so
easy to overlook the small chambers at the posterior end of the
abdomen, that the number given in some of the species may have
been under-estimated.

Pericardial Diaphragm and Space.

The heart lies in a pericardial chamber, which is bounded above by
the terga and the longitudinal tergal muscles; below by a fenestrated
membrane, the pericardial diaphragm. The intermediate space,
which is of inconsiderable depth, is nearly filled by a cellular mass
laden with fat, and resembling the fat-body.
The pericardial diaphragm, or floor of the pericardium, is continuous,
except for small oval openings scattered over its surface. It consists
of loosely interwoven fibres, interspersed with elongate nuclei
(connective-tissue corpuscles) and connected by a transparent
membrane. Into the diaphragm are inserted pairs of muscles, which,
from their shape and supposed continuity with the heart, have been
named alæ cordis, or alary muscles. 140 These are bundles of striated
muscle, about ·003 in. wide, which arise from the anterior margin of
each tergum. In the middle of the abdomen every alary muscle
passes inwards for about ·04 in., without breaking-up or widening,
and then spreads out fanwise upon the diaphragm. The fibres unite
below the heart with those of the fellow-muscle, and also join, close
to the heart, those of the muscles in front and behind. The alary
muscles are often said to distend the heart rhythmically by drawing
its walls apart, but this cannot be true. They do not pass into the
heart at all. Even if they did, a pull from opposite sides upon a
flexible, cylindrical tube, would narrow and not expand its cavity.
Moreover, direct observation 141 shows that the heart continues to
beat after all the alary muscles have been divided, and even after it
has been cut in pieces. These facts suggest that the heart of Insects
is innervated by ganglia upon or within it, and indeed transparent
larvæ, such as Corethra or Chironomus, exhibit paired cells, very like
simple ganglia, along the sides of the heart.
 Fig. 76.—Heart and Pericardial Diaphragm. On
the right, as seen from above; on the left, as
seen from below; the bottom figure rep­re­sents
a trans­verse sec­tion. Ht, heart; PD, peri­cardial
dia­phragm; AM, alary mus­cle; Tr, tra­cheal
tube; PC, peri­cardial fat-cells; PC′, multi­nuc­‐
leate fat-cells.

Scattered over the upper surface of the pericardial diaphragm are

groups of cells, similar to the fat-masses of the perivisceral space.
Over the fan-like expansions of the alary muscles are different fat-
cells, which form branched and multinucleate lobes, and radiate in
the same direction as the underlying muscles.
Tracheal trunks, arising close to the stigmata, ascend upon the
tergal wall towards the heart. They overlie the alary muscles, and
end near the heart by bifurcation, sending one branch forward and
another backward to meet corresponding branches of adjacent
trunks. A series of arches is thus formed by the dorsal tracheæ on
each side of the heart. Occasionally an arch is subdivided into two
smaller parallel tubes. A few branches of distribution are given off to
the fat-cells of the pericardium.
Graber has explained the action of the pericardial diaphragm and
chamber in the following way. 142 When the alary muscles contract,
they depress the diaphragm, which is arched upwards when at rest.
A rush of blood towards the heart is thereby set up, and the blood
streams through the perforated diaphragm into the pericardial
chamber. Here it bathes a spongy or cavernous tissue (the fat-cells),
which is largely supplied with air tubes, and having been thus
aerated, passes immediately forwards to the heart, entering it at the
moment of diastole, which is simultaneous with the sinking of the
diaphragm.
In the Cockroach the facts of structure do not altogether justify this
explanation. The fenestræ of the diaphragm are mere openings
without valves. The descent of a perforated non-valvular plate can
bring no pressure to bear upon the blood, for it is not contended
that the alary muscles are powerful enough to change the figure of
the abdominal rings. Moreover, we find comparatively few tracheal
tubes in the pericardial chamber, and can discover no proof that in
the Cockroach the fat-cells adjacent to the heart have any special
respiratory character. The diaphragm appears to give mechanical
support to the heart, resisting pressure from a distended alimentary
canal, while the sheets of fat-cells, in addition to their proper
physiological office, may equalise small local pressures, and prevent
displacement. The movement of the blood towards the heart must
(we think) depend, not upon the alary muscles, but upon the far
more powerful muscles of the abdominal wall, and upon the
pumping action of the heart itself.
 Circulation of the Cockroach.
The pulsations of the heart are rhythmical and usually frequent, the
number of beats in a given time varying with the species, the age,
and especially with the degree of activity or excitement of the Insect
observed. 143
Cornelius 144 watched the pulsations in a white Cockroach
immediately after its change of skin, and reckoned them at eighty
per minute; but he remarks that the Insect was restless, and that
the beats were probably accelerated in consequence.
In the living Insect a wave of contraction passes rapidly along the
heart from behind forwards; and the blood may under favourable
circumstances be seen to flow in a steady, backward stream along
the pericardial sinus, to enter the lateral aperture of the heart. The
peristaltic movement of the dorsal vessel may often be observed to
set in at the hinder end of the tube before the preceding wave has
reached the aorta.
From the heart a slender tube (the aorta) passes forward to the
head. It lies upon the dorsal surface of the œsophagus, which it
accompanies as far as the supra-œsophageal ganglia. In many
Insects the thoracic portion of the dorsal vessel is greatly narrowed
and non-valvular, forming the aorta of most writers on Insect
Anatomy. The aorta often dips downward near its origin, but in the
Cockroach the thoracic portion of the vessel keeps nearly the same
level as the abdominal. It gives off no lateral branches, but suddenly
ends immediately in front of the œsophageal ring in a trumpet-
shaped orifice, 145 by which the blood passes at once into a lacunar
system which occupies the perivisceral space. Here the blood bathes
the digestive and reproductive organs, receives the products of
digestion, which are not transmitted by lacteals, but discharged at
once into the blood; here, too, it gives up its urates to the excretory
tubules, and its superfluous fats to the finely-divided lobules of the
fat-body. The form of the various appendages of the alimentary
canal (salivary glands, cæcal tubes, and Malpighian tubules), as well
as of the testes, ovaries, and fat-body, is immediately connected
with the passive behaviour of the fluid upon which their nutrition
depends. Instead of being compact organs injected at every
pulsation by blood under pressure, they are diffuse, tubular, or
branched, so as to expose as large a surface as possible to the
sluggish stream in which they float.
From the perivisceral space the blood enters the pericardial sinus by
the apertures in its floor, and returns thence by the lateral inlets into
the heart.
No satisfactory injections of the circulatory channels can be made in
Insects, on account of the large lacunæ, or cavities without proper
wall, which are interposed between the heart and the extremities of
the body. In the wings and other transparent organs the blood has
been seen to flow along definite channels, which form a network,
and resemble true blood vessels in their arrangement. Whether they
possess a proper wall has not been ascertained. It is observed that
in such cases the course of the blood is generally forwards along the
anterior, and backwards along the posterior, side of the appendage.
The direction of the current is not, however, quite constant, and the
same cross branch may at different times transmit blood in different
directions. 146

Blood of the Cockroach.

The blood of the Cockroach may be collected for examination by
cutting off one of the legs, and wiping the cut end with a cover-slip.
It abounds in large corpuscles, each of which consists of a rounded
nucleus invested by protoplasm. Amœboid movements may often be
observed, and dividing corpuscles are occasionally seen. Crystals
may be obtained by evaporating a drop of the blood without
pressure; they form radiating clusters of pointed needles. The fresh-
drawn blood is slightly alkaline; it is colourless in the Cockroach, but
milky, greenish, or reddish in some other Insects. The quantity varies
greatly, according to the nutrition of the individual: after a few days’
starvation, nearly all the blood is absorbed. Larvæ contain much
more blood, in proportion to their weight, than other Insects.

Respiratory Organs of Insects.

The respiratory organs of Insects
consist of ramified tracheal tubes,
which communicate with the
external air by stigmata or spiracles.
Of these spiracles the Cockroach has
ten pairs—eight in the abdomen and
two in the thorax. The first thoracic
spiracle lies in front of the
mesothorax, beneath the edge of
the tergum; the second is similarly
placed in front of the metathorax.
The eight abdominal spiracles
belong to the first eight somites;
each lies in the fore part of its
segment, and hence, apparently, in
the interspace between two terga Fig. 77.—Tracheal System
and two sterna. The first abdominal of Cock­roach. Side view of
spiracle is distinctly dorsal in head seen from with­out,
position. intro­duc­ing the chief
branches of the left half.
The disposition of the spiracles
× 15.
observed in the Cockroach is
common in Insects, and, of all the
recorded arrangements, this approaches nearest to the plan of the
primitive respiratory system of Tracheata, in which there may be
supposed to be as many spiracles as somites. 147 The head never
carries spiracles except in Smynthurus, one of the Collembola
(Lubbock). Many larvæ possess only the first of the three possible
thoracic spiracles; in perfect Insects this is rarely or never met with
(Pulicidæ?), but either the second, or both the second and third, are
commonly developed. Of the abdominal somites, only the first eight
ever bear spiracles, and these may be reduced in burrowing or
aquatic larvæ to one pair (the eighth), while all disappear in the
aquatic larva of Ephemera.
From the spiracles, short, wide air-tubes pass inwards, and break up
into branches, which supply the walls of the body and all the viscera.
Dorsal branches ascend towards the heart on the upper side of the
alary muscles; each bifurcates above, and its divisions join those of
the preceding and succeeding segments, thus forming loops or
arches. The principal ventral branches take a transverse direction,
and are usually connected by large longitudinal trunks, which pass
along the sides of the body; the Cockroach, in addition to these,
possesses smaller longitudinal vessels, which lie close to the middle
line, on either side of the nerve-cord. 148 The ultimate branches form
an intricate network of extremely delicate tubes, which penetrates or
overlies every tissue.
Fig. 79.—Tracheal System of Cock­roach.
Back of head, seen from the front, the
fore half being re­moved. × 15. The
letters A–J indi­cate cor­res­pond­ing
branches in figs. 77, 78, and 79.
Fig. 78.—Tracheal System of Cock­roach. Top
and front of head seen from with­out. × 15.
Fig. 80.—Tracheal System of
Cock­roach. The dor­sal in­tegu­‐
ment re­moved and the vis­cera
in place. × 5.
Fig. 81.—Tracheal System of
Cock­roach. The vis­cera re­moved
to show the ven­tral tra­cheal com­‐
muni­ca­tions. × 5.
Fig. 82.—Tracheal System of
Cock­roach. The ven­tral in­‐
tegu­ment and vis­cera re­‐
moved to show the dor­sal
tra­cheal com­muni­ca­tions.
× 5.
 Fig. 83.—Tracheal tube
with its epi­the­lium and
spi­ral thread. Slight­ly
altered from a fig­ure
given by Chun (Rec­tal-
drü­sen bei den Insek­ten,
pl. iv., fig. 1).

Tracheal Tubes.
The accompanying figures sufficiently explain the chief features of
the tracheal system of the Cockroach, so far as it can be explored by
simple dissection. Leaving them to tell their own tale, we shall pass
on to the minute structure of the air-tubes, the spiracles, and the
physiology of Insect respiration.
The tracheal wall is a folding-in of the integument, and agrees with
it in general structure. Its inner lining, the intima, is chitinous, and
continuous with the outer cuticle. It is secreted by an epithelium of
nucleated, chitinogenous cells, and outside this is a thin and
homogeneous basement membrane. The integument, the tracheal
wall, and the inner layers of nearly the whole alimentary canal are
continuous and equivalent structures. The lining of the larger
tracheal tubes at least is shed at every moult, like that of the
stomodæum and proctodæum.

Tracheal Thread.
In the finest tracheal tubes (·0001 in. and
under) the intima is to all appearance
homogeneous. In wider tubes it is
strengthened by a spiral thread, which is
denser, more refractive, and more flexible
than the intervening membrane. The
thread projects slightly into the lumen of Fig. 84.—Intima
the tube, and is often branched. It is (chi­tin­ous lin­ing) of
interrupted frequently, each length making a large tra­cheal
but a few turns round the tube, and ending tube. The spir­al
in a point. The thread of a branch is never thread div­ides here
continued into a main trunk. Both the and there. Copied
thread and the intervening membrane from MacLeod, loc.
become invisible or faint when the tissue is cit., fig. 9.
soaked with a transparent fluid, so as to
expel the air. Both, but especially the thread, absorb colouring
matter with difficulty. The thread, from its greater thickness, offers a
longer resistance to solvents, such as caustic alkalies, and also to
mechanical force; it can therefore be readily unrolled, and often
projects as a loose spiral from the end of a torn tube, while the
membrane breaks up or crumbles away. 149
The large tracheal tubes close to the spiracles are without spiral
thread, and the intima is here subdivided into polygonal areas, each
of which is occupied by a reticulation of very fine threads. This
structure may be traced for a short distance between the turns of
the spiral thread.
The chitinogenous layer of the tracheal tubes is single, and consists
of polygonal, nucleated cells, forming a mosaic pattern, but
becoming irregular and even branched in the finest branches. The
cell walls are hardly to be made out without staining. Externally, the
chitinogenous cells rest upon a delicate basement membrane.
Where a number of branches are given off together, the tracheal
tube may be dilated. Fine branches, such as accompany nerves, are
often sinuous. In the very finest branches the tube loses its thread,
the chitinogenous cells become irregular, and the intima is lost in the
nucleated protoplasmic mass which replaces the regular epithelium
of the wider tubes. 150

The Spiracles.
The spiracles of the Cockroach are by no means of complicated
structure, but their small size, and the differences between one
spiracle and another, are difficulties which cost some pains to
overcome.
 Fig. 85.—First Thoracic Spiracle (left side), seen from the
out­side. × 70. V, valve; I, setose lin­ing of valve (mouth of
tra­cheal tube) × 230. The oc­clus­or mus­cle is shown. The
arrow indi­cates the direc­tion of air enter­ing the spir­acle. In
the nat­ural posi­tion this spir­acle is set oblique­ly, the slit
being inclined down­wards and back­wards. (P. ameri­cana.)

The first thoracic spiracle (fig. 85) is the largest in the body. It lies in
front of the mesothorax, between the bases of the first and second
legs. It is placed obliquely, the slit being inclined downwards and
backwards, and is closed externally by a large, slightly two-lobed
valve, attached by its lower border. The aperture immediately within
the valve divides into two nearly equal cavities, each of which leads
to a separate tracheal trunk; and between these cavities is a
septum, thickened on its free edge, against which the margin of the
valve appears to close. A special occlusor muscle arises from the
integument below the spiracle, and is inserted into a chitinous
process which projects inwardly from the centre of the valve. A
second muscle, whose connections and mode of action we have not
been able to make out satisfactorily,
lies beneath the first, and is inserted
into the thickened edge of the
septum.
The second thoracic spiracle
(fig. 86) lies in front of the
metathorax, between the bases of
the second and third legs. It is much
smaller and simpler than the first.
Its valve is nearly semi-circular, and
the free border is strengthened on
its deep surface by a chitinous rim,
which terminates beyond the end of
the hinge of the valve in a process
which gives insertion to the occlusor
muscle. Fig. 86.—Second Thoracic
Spiracle (left side), seen
The abdominal spiracles present
from the out­side. × 70. V,
quite a different plan of structure.
lower (mov­able) valve. The
The external orifice is permanently
oc­clus­or mus­cle is shown.
open, owing to the absence of
The arrow indi­cates the
valves, but communication with the
direc­tion of air enter­ing the
tracheal trunk may be cut off at
spir­acle. (P. americana.)
pleasure by an internal occluding
apparatus. The external orifice leads
into a shallow oval cup, which communicates with the tracheal trunk
by a narrow slit, or internal aperture of the spiracle. The chitinous
cuticle, surrounding this internal aperture, is richly provided with
setæ, which are turned towards the opening. 151 Fig. 87C represents
a spiracle seen from within, and shows that the slit divides the cup
into two unequal lips, the smaller of which inclines away from the
middle line of the body, is movable, and is strengthened on its deep
surface by a curved chitinous rod, the “bow” of Landois. From the
opposite lip, a pouch is thrown out, which serves for the attachment
of the occlusor muscle. The muscle is inserted into the extremity of
the bow, and when it contracts, the bow is pulled over into the
position shown in fig. 87D, and the opening is closed. The
antagonist muscle, which exists in all the abdominal spiracles, is
shown in fig. 88; it arises from the supporting plate of the spiracle,
and is inserted opposite to the occlusor, into the extremity of the
bow.
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