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Cimbex she finds a similar arrangement, but there are ten chambers,
and no aorta.

The dorsal vessel is connected with the roof of the body by some
short muscles, and is usually much surrounded by fat-body into
which tracheae penetrate; by these various means it is kept in
position, though only loosely attached; beneath it there is a delicate,
incomplete or fenestrate, membrane, delimiting a sort of space
called the pericardial chamber or sinus; connected with this
membrane are some very delicate muscles, the alary muscles,
extending inwards from the body wall (b, Fig. 72): the curtain formed
by these muscles and the fenestrate membrane is called the
pericardial diaphragm or septum. The alary muscles are not directly
connected with the heart.

Fig. 72.—Dorsal vessel (c), and alary muscles (b), of Gryllotalpa (after
Graber); a, aorta. N.B.—The ventral aspect is here dorsal, and
nearly the whole of the body is removed to show these parts.

Fig. 73.—Diagram of transverse section of pericardial sinus of

Oedipoda coerulescens. (After Graber, Arch. Mikr. Anat. ix.) H,
heart; s, septum; m, muscles—the upper suspensory, the lower
alary.

It has been thought by some that delicate vessels exist beyond the
aorta through which the fluid is distributed in definite channels, but
this does not appear to be really the case, although the fluid may
frequently be seen to move in definite lines at some distance from
the heart.

There is still much uncertainty as to some of the details of the action

of the heart, and more especially as to the influence of the alary
muscles. The effect of the contraction of these must be to increase
the area of the pericardial chamber by rendering its floor or septum
less arched, as shown in our diagram (Fig. 73), representing a
transverse section through the pericardial chamber, H being the
dorsal vessel with m its suspensory muscles, and s its septum, with
m the alary muscles. The contraction of these latter would draw the
septum into the position of the dotted line, thus increasing the area
of the sinus above; but as this floor or septum is a fenestrated
structure, its contraction allows fluid to pass through it to the
chamber above; thus this arrangement may be looked on as a
means of keeping up a supply of fluid to the dorsal vessel, the
perforated septum, when it contracts, exerting pressure on the
tissues below; these are saturated with fluid, which passes through
the apertures to the enlarged pericardial chamber.

Some misconception has prevailed, too, as to the function of the

pericardial chamber. This space frequently contains a large quantity
of fat-body—pericardial tissue—together with tracheae, and this has
given rise to the idea that it might be lung-like in function; but, as
Miall and Denny[58] have pointed out, this is erroneous; the tissues in
Insects have their own ample supplies of air. It has also been
supposed that the alary muscles cause the contraction of the heart,
but this is not directly the case, for they are not attached to it, and it
pulsates after they have been severed. It has been suggested that
the contractions of this vessel are regulated by small ganglia placed
on, or in, its substance. However this may be, these contractions
vary enormously according to the condition of the Insect; they may
be as many, it is said, as 100 or more in a minute, or they may be
very slow and feeble, if not altogether absent, without the death of
the Insect ensuing.
The expulsion of the blood from the front of the dorsal vessel seems
to be due to the rhythm of the contraction of the vessel as well as to
its mechanical structure. Bataillon says,[59] confirming an
observation of Réaumur, that at the period when the silkworm is
about to change to the chrysalis condition, the circulation undergoes
periodical changes, the fluid moving during some intervals of about
ten minutes' duration in a reversed direction, while at other times the
blood is expelled in front and backwards simultaneously, owing
apparently to a rhythmical change in the mode of contraction of the
dorsal vessel.

As the dorsal vessel consists of a number of distinct chambers, it

has been suggested that there is normally one of these for each
segment of the body; and it appears that the total number is
sometimes thirteen, which is frequently that of the segments of the
body without the head. The number of chambers differs, however,
greatly, as we have previously stated, and cannot be considered to
support the idea of an original segmental arrangement of the
chambers. The dorsal vessel, though in the adult a single organ,
arises in the embryo from two lateral, widely separated parts which
only in a subsequent stage of the embryonic development coalesce
in the median line.

Fat-Body.

In discussing the tracheae we remarked on the importance of their

function and on their abundant presence in the body. Equally
conspicuous, and perhaps scarcely less important in function, is the
fat-body, which on opening some Insects, especially such as are in
the larval stage, at once attracts attention. It consists of masses of
various size and indefinite form distributed throughout the body,
loosely connected together, and more or less surrounding and
concealing the different organs. The colour varies according to the
species of Insect. This fat-body is much connected with fine tracheal
twigs, so that an organisation extending throughout the body is thus
formed. It may be looked on as a store of nutritious matter which
may be added to or drawn on with great rapidity; and it is no doubt
on this that many of the internal parasites, so common in the earlier
stages of Insects' lives, subsist before attacking the more permanent
tissues of their hosts. There is some reason to suppose that the fat-
body may have some potency in determining the hunger of the
Insect, for some parasitised larvae eat incessantly.

The matter extracted from the food taken into the stomach of the
Insect, after undergoing some elaboration—on which point very little
is known—finds its way into the body-cavity of the creature, and as it
is not confined in any special vessels the fat-body has as unlimited a
supply of the nutritive fluid as the other organs: if nutriment be
present in much greater quantity than is required for the purposes of
immediate activity, metamorphosis or reproduction, it is no doubt
taken up by the fat-body which thus maintains, as it were, an
independent feeble life, subject to the demands of the higher parts of
the organisation. It undoubtedly is very important in metamorphosis,
indeed it is possible that one of the advantages of the larval state
may be found in the fact that it facilitates, by means of the fat-body,
the storage in the organisation of large quantities of material in a
comparatively short period of time.

A considerable quantity of fat tissue is found in the pericardial sinus,

where it is frequently of somewhat peculiar form, and is spoken of as
pericardial cells, or pericardial tissue. Some large cells, frequently of
pale yellow colour, and containing no fat, are called oenocytes by
Wielowiejski. They are connected with the general fat-body, but are
not entirely mingled with it; several kinds have been already
distinguished, and they are probably generally present. The
phagocytes, or leucocytes, the cells that institute the process of
histolysis in the metamorphosis of Muscidae, are a form of blood
cell; though these cells are amoeboid some writers derive them from
the fat-body. The cells in the blood have no doubt generally an
intimate relation with the fat-body, but very little accurate information
has been obtained as to these important physiological points, though
Graber has inaugurated their study.[60]

Organs of Sex.

The continuation of the species is effected in Insects by means of

two sexes, each endowed with special reproductive organs. It has
been stated that there are three sexes in some Insects—male,
female, and neuter; but this is not correct, as the so-called neuters
are truly sexed individuals,—generally females,—though, as a rule,
they are not occupied with the direct physiological processes for
continuing the species.

The offspring is usually produced in the shape of eggs, which are

formed in ovaries. These organs consist of egg-tubes, a cluster of
which is placed on each side of the body, and is suspended,
according to Leydig[61] and others, to the tissue connected with the
heart by means of the thread-like terminations of the tubes.

Fig. 74.—Sex organs of female of Scolia interrupta (after Dufour); a,

egg-tubes; b, oviducts; c, poison glands; d, duct of accessory
gland (or spermatheca); e, external terminal parts of body.

The number of egg-tubes varies greatly in different Insects; there

may be only one to each ovary (Campodea), but usually the number
is greater, and in the queen-bee it is increased to about 180. In the
Queens of the Termitidae, or white ants, the ovaries take on an
extraordinary development; they fill the whole of the greatly
distended hind-body. Three thousand egg-tubes, each containing
many hundred eggs, may be found in a Queen Termite, so that, as
has been said by Hagen,[62] an offspring of millions in number is
probable. There is considerable variety in the arrangements for the
growth of the eggs in the egg-tubes. Speaking concisely, the tubes
may be considered to be centres of attraction for nutritive material, of
which they frequently contain considerable stores. Next to the
terminal thread, of which we have already spoken, there is a greater
or smaller enlargement of the tube, called the terminal chamber; and
there may also be nutriment chambers, in addition to the dilatations
which form the egg-chambers proper. Korschelt[63] distinguishes
three principal forms of egg-tubes, viz. (1) there are no special
nutriment chambers, a condition shown in Figure 74; (2) nutriment
chambers alternate with the egg-chambers, as shown in our Figure
of an egg-tube of Dytiscus marginalis; (3) the terminal chamber
takes on an unusual development, acting as a large nutriment
chamber, there being no other special nutriment chambers. This
condition is found in Rhizotrogus solstitialis. The arrangements as to
successive or simultaneous production of the eggs in the tubes
seem to differ in different Insects. In some forms, such as the white
ants, the process of egg-formation (oogenesis) attains a rapidity that
is almost incredible, and is continued, it is said, for periods of many
months. There is no point in which Insects differ more than in that of
the number of eggs produced by one female. The egg-tubes are
connected with a duct for the conveyance of the eggs to the exterior,
and the arrangements of the tubes with regard to the oviduct also
vary much. An interesting condition is found in Machilis (see Fig. 94,
p. 188), where the seven egg-tubes are not arranged in a bunch, but
open at a distance from one another into the elongated duct. The
two oviducts usually unite into one chamber, called the azygos
portion or the uterus, near their termination. There are a few Insects
(Ephemeridae) in which the two oviducts do not unite, but have a
pair of orifices at the extremity of the body. Hatchett-Jackson has
recently shown[64] that in Vanessa io of the Order Lepidoptera, the
paired larval oviducts are solid, and are fixed ventrally so as to
represent an Ephemeridean stage; that the azygos system of ducts
and appended structures develop separately from the original
oviducts, and that they pass through stages represented in other
Orders of Insects to the stage peculiar to the Lepidoptera. Machilis,
according to Oudemans, is a complete connecting link between the
Insects with single and those with paired orifices.

There are in different Insects more than one kind of diverticula and
accessory glands in connexion with the oviducts or uterus; a
receptaculum seminis, also called spermatheca, is common. In the
Lepidoptera there is added a remarkable structure, the bursa
copulatrix, which is a pouch connected by a tubular isthmus with the
common portion of the oviduct, but having at the same time a
separate external orifice, so that there are two sexual orifices, the
opening of the bursa copulatrix being the lower or more anterior. The
organ called by Dufour in his various contributions glande sébifique,
is now considered to be, in some cases at any rate, a spermatheca.
The special functions of the accessory glands are still very obscure.

Fig. 75.—Egg-tube of Dytiscus marginalis; e.c, egg-chamber; n.c,

nutriment chamber; t.c, terminal chamber; t.t, terminal thread.
(After Korschelt.)

The ovaries of the female are replaced in the male by a pair of

testes, organs exhibiting much variety of form. The structure may
consist of an extremely long and fine convoluted tube, packed into a
small space and covered with a capsule; or there may be several
shorter tubes. As another extreme may be mentioned the existence
of a number of small follicles opening into a common tube, several of
these small bodies forming together a testis. As a rule each testis
has its own capsule, but cases occur—very frequently in the
Lepidoptera—in which the two testes are enclosed in a common
capsule; so that there then appears to be only one testis. The
secretion of each testis is conveyed outwards by means of a slender
tube, the vas deferens, and there are always two such tubes, even
when the two testes are placed in one capsule. The vasa deferentia
differ greatly in their length in different Insects, and are in some
cases many times the length of the body; they open into a common
duct, the ductus ejaculatorius. Usually at some part of the vas
deferens there exists a reservoir in the form of a sac or dilatation,
called the vesicula seminalis. There are in the male, as well as in the
female, frequently diverticula, or glands, in connexion with the sexual
passages; these sometimes exhibit very remarkable forms, as in the
common cockroach, but their functions are quite obscure. There is,
as we have already remarked, extreme variety in the details of the
structure of the internal reproductive apparatus in the male, and
there are a few cases in which the vasa deferentia do not unite
behind, but terminate in a pair of separate orifices. The genus
Machilis is as remarkable in the form of the sexual glands and ducts
of the male as we have already mentioned it to be in the
corresponding parts of the female.

Fig. 76.—Tenthredo cincta. a, a, testes; b, b, vasa deferentia; c, c,

vesiculæ seminales; d, extremity of body with copulatory
armature. (After Dufour.)

Although the internal sexual organs are only fully developed in the
imago or terminal stage of the individual life, yet in reality their
rudiments appear very early, and may be detected from the embryo
state onwards through the other preparatory stages.

The spermatozoa of a considerable number of Insects, especially of

Coleoptera, have been examined by Ballowitz;[65] they exhibit great
variety; usually they are of extremely elongate form, thread-like, with
curious sagittate or simply pointed heads, and are of a fibrillar
structure, breaking up at various parts into finer threads.

External Sexual Organs.—The terminal segments of the body are

usually very highly modified in connexion with the external sexual
organs, and this modification occurs in such a great variety of forms
as to render it impossible to give any general account thereof, or of
the organs themselves. Some of these segments—or parts of the
segments, for it may be dorsal plates or ventral plates, or both—may
be withdrawn into the interior, and changed in shape, or may be
doubled over, so that the true termination of the body may be
concealed. The comparative anatomy of all these parts is especially
complex in the males, and has been as yet but little elucidated, and
as the various terms made use of by descriptive entomologists are of
an unsatisfactory nature we may be excused from enumerating
them. We may, however, mention that when a terminal chamber is
found, with which both the alimentary canal and the sexual organs
are connected, it is called a cloaca, as in other animals.

Parthenogenesis.

There are undoubted cases in Insects of the occurrence of

parthenogenesis, that is, the production of young by a female without
concurrence of a male. This phenomenon is usually limited to a
small number of generations, as in the case of the Aphididae, or
even to a single generation, as occurs in the alternation of
generations of many Cynipidae, a parthenogenetic alternating with a
sexual generation. There are, however, a few species of Insects of
which no male is known (in Tenthredinidae, Cynipidae, Coccidae),
and these must be looked on as perpetually parthenogenetic. It is a
curious fact that the result of parthenogenesis in some species is the
production of only one sex, which in some Insects is female, in
others male; the phenomenon in the former case is called by
Taschenberg[66] Thelyotoky, in the latter case Arrhenotoky;
Deuterotoky being applied to the cases in which two sexes are
produced. In some forms of parthenogenesis the young are
produced alive instead of in the form of eggs. A very rare kind of
parthenogenesis, called paedogenesis, has been found to exist in
two or three species of Diptera, young being produced by the
immature Insect, either larva or pupa.

Glands.

Insects are provided with a variety of glands, some of which we have

alluded to in describing the alimentary canal and the organs of sex;
but in addition to these there are others in connexion with the outer
integument; they may be either single cells, as described by Miall in
Dicranota larva,[67] or groups of cells, isolated in tubes, or pouches.
The minute structure of Insect glands has been to some extent
described by Leydig;[68] they appear to be essentially of a simple
nature, but their special functions are very problematic, it being
difficult to obtain sufficient of their products for satisfactory
examination.

CHAPTER V

DEVELOPMENT

EMBRYOLOGY–EGGS–MICROPYLES–FORMATION OF EMBRYO–VENTRAL
PLATE–ECTODERM AND ENDODERM–SEGMENTATION–LATER STAGES–
DIRECT OBSERVATION OF EMBRYO–METAMORPHOSIS–COMPLETE AND
 INCOMPLETE–INSTAR–HYPERMETAMORPHOSIS–METAMORPHOSIS OF
INTERNAL ORGANS–INTEGUMENT–METAMORPHOSIS OF BLOWFLY–
HISTOLYSIS–IMAGINAL DISCS–PHYSIOLOGY OF METAMORPHOSIS–
ECDYSIS.

The processes for the maintenance of the life of the individual are in
Insects of less proportional importance in comparison with those for
the maintenance of the species than they are in Vertebrates. The
generations of Insects are numerous, and the individuals produced
in each generation are still more profuse. The individuals have as a
rule only a short life; several successive generations may indeed
make their appearances and disappear in the course of a single
year.

Although eggs are laid by the great majority of Insects, a few species
nevertheless increase their numbers by the production of living
young, in a shape more or less closely similar to that of the parent.
This is well known to take place in the Aphididae or green-fly Insects,
whose rapid increase in numbers is such a plague to the farmer and
gardener. These and some other cases are, however, exceptional,
and only emphasise the fact that Insects are pre-eminently
oviparous. Leydig, indeed, has found in the same Aphis, and even in
the same ovary, an egg-tube producing eggs while a neighbouring
tube is producing viviparous individuals.[69] In the Diptera pupipara
the young are produced one at a time, and are born in the pupal
stage of their development, the earlier larval state being undergone
in the body of the parent: thus a single large egg is laid, which is
really a pupa.

The eggs are usually of rather large size in comparison with the
parent, and are produced in numbers varying according to the
species from a few—15 or even less in some fossorial Hymenoptera
—to many thousands in the social Insects: somewhere between 50
and 100 may perhaps be taken as an average number for one
female to produce. The whole number is frequently deposited with
rapidity, and the parent then dies at once. Some of the migratory
locusts are known to deposit batches of eggs after considerable
intervals of time and change of locality. The social Insects present
extraordinary anomalies as to the production of the eggs and the
prolongation of the life of the female parent, who is in such cases
called a queen.

The living matter contained in the egg of an Insect is protected by

three external coats: (1) a delicate interior oolemm; (2) a stronger,
usually shell-like, covering called the chorion; (3) a layer of material
added to the exterior of the egg from glands, at or near the time
when it is deposited, and of very various character, sometimes
forming a coat on each egg and sometimes a common covering or
capsule for a number of eggs. The egg-shell proper, or chorion, is
frequently covered in whole or part with a complex minute sculpture,
of a symmetrical character, and in some cases this is very highly
developed, forming an ornamentation of much delicacy; hence some
Insects' eggs are objects of admirable appearance, though the
microscope is of course necessary to reveal their charms. One of the
families of butterflies, the Lycaenidae, is remarkable for the complex
forms displayed by the ornamentation of the chorion (see Fig. 78, B).

Fig. 77.—Upper or micropylar aspect of egg of Vanessa cardui. (After

Scudder.)

The egg-shell at one pole of the egg is perforated by one or more

minute orifices for the admission to the interior of the spermatozoon,
and it is the rule that the shell hereabouts is symmetrically sculptured
(see Fig. 77), even when it is unornamented elsewhere: the
apertures in question are called micropyles. They are sometimes
protected by a micropyle apparatus, consisting of raised processes,
or porches: these are developed to an extraordinary extent in some
eggs, especially in those of Hemiptera-Heteroptera (see Fig. 78, C).
Some of these peculiar structures have been described and figured
by Leuckart.[70] The purpose they serve is quite obscure.

Fig. 78.—Eggs of Insects: A, blowfly (after Henking); B, butterfly,

Thecla (after Scudder); C, Hemipteron (Reduviid).

Formation of Embryo.

The mature, but unfertilised, egg is filled with matter that should
ultimately become the future individual, and in the process of
attaining this end is the seat of a most remarkable series of changes,
which in some Insects are passed through with extreme rapidity. The
egg-contents consist of a comparatively structureless matrix of a
protoplasmic nature and of yolk, both of which are distributed
throughout the egg in an approximately even manner. The yolk,
however, is by no means of a simple nature, but consists, even in a
single egg, of two or three kinds of spherular or granular
constituents; and these vary much in their appearance and
arrangement in the early stages of the development of an egg, the
yolk of the same egg being either of a homogeneously granular
nature, or consisting of granules and larger masses, as well as of
particles of fatty matter; these latter when seen through the
microscope looking sometimes like shining, nearly colourless,
globules.
 Fig. 79.—Showing the two extruded polar bodies P1, P2 now nearly
fused and reincluded, and the formation of the spindle by junction
of the male and female pronuclei. (After Henking.)

The nature of the matrix—which term we may apply to both the

protoplasm and yolk as distinguished from the minute formative
portions of the egg—and the changes that take place in it have been
to some extent studied, and Kowalewsky, Dohrn,[71] Woodworth,[72]
and others have given some particulars about them. The early
changes in the formative parts of the mature egg have been
observed by Henking in several Insects, and particularly in
Pyrrhocoris, his observations being of considerable interest. When
the egg is in the ovary and before it is quite mature,—at the time, in
fact, when it is receiving nutriment from ovarian cells,—it contains a
germinal vesicle including a germinal spot, but when the egg is
mature the germinal vesicle has disappeared, and there exists in its
place at one portion of the periphery of the egg-contents a cluster of
minute bodies called chromosomes by Henking, whom we shall
follow in briefly describing their changes. The group divides into two,
each of which is arranged in a rod or spindle-like manner, and may
then be called a directive rod or spindle. The outer of these two
groups travels quite to the periphery of the egg, and there with some
adjacent matter is extruded quite outside the egg-contents (not
outside the egg-coverings), being in its augmented form called a
polar or directive body. While this is going on the second directive
spindle itself divides into two groups, the outer of which is then
extruded in the manner we have already described in the case of the
first polar body, thus completing the extrusion of two directive bodies.
The essential parts of the bodies that are successively formed during
these processes are the aggregates, called chromosomes; the
number of these chromosomes appears to be constant in each
species; their movements and dispositions are of a very interesting
character, the systems they form in the course of their development
having polar and equatorial arrangements. These we cannot further
allude to, but may mention that the extrusion of the directive bodies
is only temporary, they being again included within the periphery of
the egg by the growth and extension of adjacent parts which meet
over and thus enclose the bodies.

The arrangements and movements we have briefly alluded to have

been limited to the unfertilised condition of the egg (we should rather
say, the fertilising element has taken no part in them), and have as
their result the union of the chromosomes existing after the extrusion
of the two polar bodies, into a small body called the female
pronucleus or egg-nucleus (Eikern), while the position of the
movements has been an extremely minute portion of the egg near to
its outer surface or periphery. The introduction of a sperm, or male,
element to the egg through the micropyle gives rise to the formation
of another minute body placed more in the interior of the egg, and
called the sperm-nucleus. The egg-nucleus, travelling more into the
interior of the egg, meets the sperm-nucleus; the two amalgamate,
forming a nucleus or body that goes through a series of changes
resulting in its division into two daughter-bodies. These two again
divide, and by repetitions of such division a large number of nuclei
are formed which become arranged in a continuous manner so as to
form an envelope enclosing a considerable part (if not quite the
whole) of the egg-mass. This envelope is called the blastoderm, and
together with its contents will form the embryo. We must merely
allude to the fact that it has been considered that some of the nuclei
forming the blastoderm arise directly from the egg-mass by a
process of amalgamation, and if this prove to be correct it may be
admitted that some portions of the embryo are not entirely the result
of division or segmentation of combined germ and sperm-nuclei.
Wheeler states[73] that some of the nuclei formed by the first
differentiation go to form the vitellophags scattered throughout the
yolk. We should also remark that, according to Henking, the
blastoderm when completed shows at one part a thickening,
immediately under which (i.e. included in the area the blastoderm
encloses) are the two polar bodies, which, as we have seen, were
formed by the germinating body at an earlier stage of its activity. Fig.
79 represents a stage in the development of Pyrrhocoris, showing
the interior of the egg after a body has been formed by the union of
the sperm and egg-nuclei; this body is about to undergo division or
segmentation, and the equatorial arrangement where this will take
place is seen. The two polar bodies P1, P2, after having been
excluded, are nearly reincluded in the egg.

The Ventral Plate.

The next important change after the formation of the blastoderm is

the partial detachment of a part of its periphery to become placed in
the interior of the other and larger portion. The way in which this
takes place will be gathered from the accompanying diagrammatic
figures taken from Graber: a thickened portion (a b) of the
blastoderm becomes indrawn so as to leave a fold (c d) at each point
of its withdrawal, and these folds afterwards grow and meet so as to
enclose the thickened portion. The outer envelope, formed in part by
the original blastoderm and in part by the new growth, is called the
serosa (e f), the inner layer (g) of the conjoined new folds being
termed the amnion: the part withdrawn to the interior and covered by
the serosa and amnion is called the ventral plate, or germinal band
(Keimstreif), and becomes developed into the future animal. The
details of the withdrawal of the ventral plate to the interior are very
different in the various Insects that have been investigated.
 Fig. 80.—Stages of the enclosure of the ventral plate: A, a, b, ventral plate;
B, c, d, folds of the blastoderm that form the commencement of the
amnion and serosa; C, e, f, part of the serosa; g, amnion.

One of the earliest stages in the development is a differentiation of a

portion of the ventral plate into layers from which the future parts of the
organisation will be derived. This separation of endoderm from ectoderm
takes place by a sort of invagination, analogous with that by which the
ventral plate itself is formed. A longitudinal depression running along the
middle of the ventral plate appears, and forms a groove or channel, which
becomes obliterated as to its outer face by the meeting together of the
two margins of the groove (except on the anterior part, which remains
open). The more internal layer of the periphery of this closed canal is the
origin of the endoderm and its derivatives. Subsequently the ventral plate
and its derivatives grow so as to form the ventral part and the internal
organs of the Insect, the dorsal part being completed much later by
growths that differ much in different Insects; Graber, who has specially
investigated this matter, informing us[74] that an astonishing
multifariousness is displayed. It would appear that the various modes of
this development do not coincide with the divisions into Orders and
Families adopted by any systematists.

We should observe that the terms ectoderm, mesoderm, and endoderm

will probably be no longer applied to the layers of the embryo when
embryologists shall have decided as to the nature of the derived layers,
and shall have agreed as to names for them. According to the
nomenclature of Graber[75] the blastoderm differentiates into Ectoblast
and Endoblast; this latter undergoing a further differentiation into
Coeloblast and Myoblast. This talented embryologist gives the following
table of the relations of the embryonic layers and their nomenclature, the
first term of each group being the one he proposed to use:—
Nussbaum considers[76] that "there are four layers in the cockroach-
embryo, viz. (1) epiblast, from which the integument and nervous system
are developed; (2) somatic layer of mesoblast, mainly converted into the
muscles of the body-wall; (3) splanchnic layer of mesoblast, yielding the
muscular coat of the alimentary canal; and (4) hypoblast, yielding the
epithelium of the mesenteron."

Fig. 81.—Early stages of the segmentation of a beetle (Lina): A,

segmentation not visible, 1 day; B, segmentation of head visible; C,
segmentation still more advanced, 2¼ days; PC, procephalic lobes; g1,
g2, g3, segments bearing appendages of the head; th, thorax; th1, th2,
th3, segments of the thorax; a1, a2, anterior abdominal.

Turning our attention to the origin of the segmentation, that is so marked

a feature of Insect structure, we find that evidence of division or
arrangement of the body into segments appears very early, as shown in
our Figure of some of the early stages of development of Lina (a beetle),
Fig. 81. In A the segmentation of the ectoderm has not commenced, but
the procephalic lobes (P C) are seen; in B the three head segments are
distinct, while in C the thoracic segmentation has occurred, and that of
the abdomen has commenced. Graber considers that in this species the
abdomen consists of ten segmental lobes, and a terminal piece or telson.
According to Graber[77] this is not a primitive condition, but is preceded by
a division into three or four parts, corresponding with the divisions that will
afterwards be head, thorax, and abdomen. This primary segmentation, he
says, takes place in the Hypoblast (Endoderm) layer of the ventral plate;
this layer being, in an early stage of the development of a common
grasshopper (Stenobothrus variabilis), divided into four sections, two of
which go to form the head, while the others become thorax and abdomen
respectively. In Lina the primary segmentation is, Graber says, into three
instead of four parts. Graber's opinion on the primary segmentation does
not appear to be generally accepted, and Wheeler, who has studied[78]
the embryology of another Orthopteron, considers it will prove to be
incorrect. When the secondary segmentation occurs the anterior of the
two cephalic divisions remains intact, while the second divides into the
three parts that afterwards bear the mouth parts as appendages. The
thoracic mass subsequently segments into three parts, and still later the
hind part of the ventral plate undergoes a similar differentiation so as to
form the abdominal segments; what the exact number of these may be is,
however, by no means easy to decide, the division being but vague,
especially posteriorly, and not occurring all at once, but progressing from
before backwards.

The investigations that have been made in reference to the segmentation

of the ventral plate do not at present justify us in asserting that all Insects
are formed from the same number of embryonic segments. The matter is
summarised by Lowne, to the effect that posterior to the procephalic lobes
there are three head segments and three thoracic segments, and a
number of abdominal segments, "rarely less than nine or more than
eleven." It will be seen by referring to Figure 81 that the segmentation
appears, not simultaneously, but progressively from the head backwards;
this of course greatly increases the difficulty of determining by means of a
section the real number of segments.
 Fig. 82.—Embryo of a moth (Zygaena) at the fifth day (after Graber): am,
amnion; s, serosa; p, procephalic lobes; st, stomodaeum; pr,
proctodaeum; g1, g2, g3, the mouth parts or head appendages; th1, th2,
th3, appendages of the thoracic segments; a1-a10, abdominal
segments; s.g, salivary gland.

The later stages in the development of Insects are already proved to be

so various that it would be impossible to attempt to follow them in detail;
but in Fig. 82 we represent a median section of the embryo of Zygaena
filipendula at the fifth day. It shows well some of the more important of the
general features of the development at a stage subsequent to those
represented in Fig. 81, A, B, C. The very distinct stomodaeum (st) and
proctodaeum (pr) are seen as inflexions of the external wall of the body;
the segmentation and the development of the ventral parts of the embryo
are well advanced, while the dorsal part of the embryo is still quite
incomplete.

The method of investigation by which embryologists chiefly carry on their

researches is that of dividing the egg after proper preparation, into a large
number of thin sections, which are afterwards examined in detail, so as to
allow the arrangement to be completely inferred and described. Valuable
as this method is, it is nevertheless clear that it should, if possible, be
supplemented by direct observation of the processes as they take place
in the living egg: this method was formerly used, and by its aid we may
still hope to obtain exact knowledge as to the arrangements and
rearrangements of particles by which the structures develop. Such
questions as whether the whole formative power in the egg is absolutely
confined to one or two small centres to which the whole of the other egg
contents are merely, as it were, passive accessories, or whether an egg is
a combination in which some portion of the powers of rearrangement is
possessed by other particles, as well as the chromosomes, in virtue of
their own nature or of their position at an early period in the whole, can
scarcely be settled without the aid of direct observation of the processes
during life.

The importance of the yolk is recognised by most of the recent writers.

Nussbaum states (loc. cit.) that "scattered yolk-cells associate themselves
with the mesoblast cells, so that the constituents of the mesoblast have a
twofold origin." Wheeler finds[79] that amoeboid cells—he styles them