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Structure, function and taxonomic value of the proventriculus of mandibulate insects

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STHJCTURE, FUNCTION AMD TAXONOMIC VAIIJE
of
THE PROVSNTBICULUS
of
2D
MAM.1BUIA 'IE INSECTS
By
W. 1. JUDD.
Thesis presented in partial fulfilment
for
the degree of Master of Arts
at
The University of Western Ontario
May 1940,
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UMI Number: EC54042
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ACOQW LBDGrEMENTS
Special acknowledgement is due to Dr. J. D. Detwiler,
Professor of Applied Biology, under whose guidance the
study was made, for his interest and advice,
I am much in­
debted to Professor E. M. Walker of the University of foronto
Dr. T. H. Frison, Chief of the Natural History Survey, Urbana
Illinois, and Mr. W. J. Brown, Entomologist, Department of
Agriculture, Ottawa for identification of specimens.
I also
wish to thank Dr. A. W. A. Brown, Entomologist in charge of
the Forest Insect Survey for supplying insect specimens, and
Mr. S. Calvert of the Forest Insect Survey for aid in photo­
graphic work.
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TABLE OF COITSTTS
Page.
INTRODUCTION............................................. 1
MORPHOLOGY
............................................. 2,
Th.ysan.ura
.
................................
Collemhola
4,
......................................5
Plecoptera........................................... 6,
Neuroptera
Odonata
.
................................ 7,
..........
7,
Meooptera........................................... 8.
Bphemerida . . . . . . . . . .
Thysanoptera . . . . .
Triehoptera
........................ 8.
............................... 8.
................................
8.
Lepidoptera....................
10,
Coleoptera
..................
10,
Dermaptera
..........
19.
Orthoptera
............................... 20.
Hymenoptera........................................ 28.
CORRELATION OF THE STRUCTURE OF THE PROVSiTRICULUS WITH
FEEDING HABITS
..................................... 31.
Literature..................
.31,
Disoussion
.31.
.........
Carnivorous Predators............................... 36,
Herbivores..............
37,
Feeders on hardwood............
37,
Sucking Insects
Non-feeders
.
................................. 38,
. . .
................................. 38,
Continuous feeders on solid foods
............... 38,
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FUNCTION OF THE PEOVMTRICULUS............
.39.
Literature
39.
Funotion of the proventriculus of Gryllus
domestious
Latr.
......
..................
Funotion of the proventriculus in Acridiidae
......
40*
43.
TAXONOMIC VALUE OF THE PBQVEBITRICUUJS..................... 43.
Literature
........
....................
. . . 43.
Key for separation of four genera of Gryllidae........ 45.
S U M M A H f ................................................ 47.
MATERIALS AND M E T H O D S ....................................48.
Insects collected
................................. 48.
Histological technique..................
Whole Mounts
Photographs
LITERATURE CITED
51.
.................................... 51.
....................................... 52.
......................................
PLATES.............................
53.
69.
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liTRODUGTIOH
The proventriculus, as the term implies is the region of the fore-gut
which lies immediately in front of the ventriculus or the mid-gut of insects.
In oommon with the rest of the fore-gut it is ectodermal in origin, being
lined with a chitinous intima, in contrast to the mid-gut which is endoder­
mal in origin and which has no chitinous intima.
As will he shown in subse­
quent descriptions, the proventriculus displays all degrees of development
from a simple sphincter lined with soft cuticle to a powerful muscular organ
armed with spines and teeth*
Insects are not the only Arthropods which have the fore-gut and mid-gut
arising respectively from ectodermal and endodermal origin.
In Peripatopsis,
of the Onyohophora, this condition is found (67), as also in Briophyes tristriatus of the Acarina (66) and in Myriapoda (102,105).
While the term proventriculus is properly applied only to the hind region
of the fore-gut, some authors use it with reference to a swollen region of the
mid-gut of Diptera.
Snodgrass (122) shows that this is not truly a proven­
triculus since it is not ectodermal in origin and since the oesophageal valve
projects into its anterior end, rather than from its posterior end into the
mid-gut.
The work of several authors on Diptera confirms this view (10, 22,
37, 64, 65, 76, 78, 130, 138).
In some Diptera notably in Drosophila mel-
anogaster, the "proventricuius” is composed of both ectodermal and endoder­
mal tissues (108, 136).
Among the Homoptera, the region desoribed as pro­
ventriculus is also seen to be of endodermal origins
didae (72, 73).
Coocidae (79), Cioo-
In the Heteroptera the whole fore-gut is reduced in size
(25,B3) although the oesophageal valve is well defined in some cases.
some predatory Henriptera, there is a distinct proventriculus (47).
In
It is
thus to be seen that a specialized proventriculus in the fore-gut of sucking
—
1
—
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insects is rare*
2 -
She most important exception to this generalization is to
he seen in the Siphonaptera of which several species have a globular proven­
triculus lined with spines (22,51,95,135).
It is in mandibulate insects that the proventriculus shows the widest
variation in structure.
At the same time there are some oonstant characters!
a chitinous intima secreted by a layer of epithelial cells, and layers of
longitudinal and circular muscle.
At its posterior end, the proventriculus
projects into the mid-gut as an oesophageal v&lve.
Wigglesworth
It is maintained by
(134) that this valve secretes the peritrophic membrane and
presses it into shape, and Brown (26) suggests that this membrane contains a
small amount of chitin.
The great diversity of structure shown by the proventriculus has led to
investigations along three main avenues:
1.
A study of the correlation between the structure of the proven­
triculus and the feeding habits of insects.
2.
A study of the function of the proventriculus, and
3.
A study of hie proventriculusas a structure to be used in
classification of insects as also in investigation of phylogenetic re­
lationships.
Prom a general review of the literature it is the writer's opinion that
the interpretation of the relationship between structure of the proventricu­
lus and feeding habits of insects is still unsatisfactory.
clear up some of the difficulties
the mandibulated orders.
It is proposed to
by a study of representatives of most of
This will be enlarged and supplemented by consider­
ation of forms studied by several authors.
MOHPHOLOGY
Insects which were dissected and studied in the laboratory are the fol­
lowing!
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Thysanura:
Machilis sp.
Plecoptera:
Aoroneuria abnormis Hewm.
Trichopterat
Macroneraa zebra-bum Hagen
Coleopteras
Cicindelidae:
Carabidaet
Meloidaej
Cioindela sex-guttata Fab.
Chlaenius sp., Brachyma sp.
Macrobasis unioolor Kirby.
Stapbylinidae:
Elateridaes
Staphylinus badipes Lee.
Ludius appropinquans.
Curculionidae:
Hypomolyx piceus Be G.
Ortbopteras
Phasmidae:
Mantidae:
Acridudae:
Diapheromera femorata Say
Mantis religiosa L.
Dissoateira Carolina L.
Encoptolophus sordidus Burn.
Melanoplus femur-rubrum Be G.
M. bivittatus Say
M. punctulatus Scudd.
Chorthippus curtipennis Harris
fettigoniidaes
Conocephalus fasciatus De G.
C. brevipennia Scudd.
Neoconocephalus ensiger Harris
Soudderia curvicauda De G.
Ceuthophilus maculatus Harris
Gryllidae:
Oecanthus nigricomis. Walker
0. niveus De G.
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Gryllua pemsylvanicus Buna.
G. domesticus Latr.
Uemobius fasciatus De G.
Gryllotalpa sp#
Hymenoptera:
Tenthridinidaej
larvae of Meodiprion lecontei Pitch, Diprion polytomum
Htg., Arge peotoralis Leach, Cephalcia Sp.
Information on the structure of many other species of insects was found hy
reviewing literature on the digestive tracts of insects.
Such references are
referred to hy numbers in brackets corresponding to the numbers of publications
listed in the bibliography.
fhysanura
Machilis sp.: In Machilis the oesophagus occupies the prothorax and mesothorax
(PI. 1. Pig. 1-oes.} In the fore part of the mesothorax it projects into
the mid-gut, forming the oesophogeal valve.
In the prothorax the oesopha­
gus is 130 M. in diameter and is surrounded by a single band of circular mus­
cle, 3-4 M. in -thickness (PI. 1. Pig. 2-CM). The epithelial layer is made up
of a single layer of cells roughly cuboidal in shape and 5-9 M. in thickness
(JSP).
The chitinous intima (1) is composed of two layers, an outer one
ranging from 1-10 M. in thickness and an inner one with a constant thickness
of 1 M.
The intima is thrown up into longitudinal ridges of irregular shape#
In the posterior part of the prothorax the oesophagus has a diameter of about
140
M. and the sheath of circular muscle which surrounds it is 2 M. thick.
(PI. 1.
Pig. B). The ridges of the intima in the lateral walls of the oeso­
phagus are flattened out, utile dorsally and ventrally the intima projects in-
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ward forming dorsal and ventral ridges (PI* 1 Fig. 3 -DB,Vfi).
the lumen between these ridges is about 5 M.
The width of
In the fore part of the meso­
thorax, the posterior end of the oesophagus forms the oesophageal valve. (PI.
1 Fig. 4-OES?).
the mid-gut.
It is surrounded by two lobes (L) which project forward from
The vertical diameter of the valve is about 180 M, the transverse
diameter about 90 M.
In this region the dorsal and ventral ridges of the oeso­
phagus touch one another and are broadened out laterally to touch the lateral
walls (PI, 1 Fig. 5).
In this way the lumen is much reduced.
A cross-section
of this region (Fig. 5} shows on the outside the irregular layer of intima (lij
which is folded back to join the cells of the mid-gut (MG).
Inside this layer
are two layers of epithelium (SP) separated by narrow strands of circular mus­
cle (CM).
Inside the inner layer of epithelium is the chitinous intima of the
valve (II).
Lepiama has a more specialized proventricular region than has Machilis*
Jfewcomer (92) reports that it has a muscular crop, and Gray (6) points out
that its proventriculus is spherical and is provided with six teeth.
Collembola
MacNamara (83) studied the food habits of Collembola and showed that
some of the species are carnivorous, but that most species are vegetarian,
feeding on molds, algae and decayed tissues.
The structure of the digestive
canal has been studied by several investigators.
Sommer (123) studied a species
of family Poduridae and found that its pro-ventricular region is similar to
that of Machilis. In Sine11a coeca and Tomooerus vulgaris
Boelitz (11) found
a proventricular region which also resembles that of Machilis. There is a thin
layer of circular muscle, a narrow lining of ohitin and dorsal and ventral
ridges projecting into the lumen.
A similar condition is shown by Mukerji (88)
to exist in Protanura oarpenteri sp. now.
In this form the fore-gut projects
into the mid -gut as a short oesophageal valve*
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Pleooptera
Perlidaes
Acroneurla abnormia lewm.: In the larva of this inseot the pro­
ventriculus is cylindrical and when the larva is fully grown it is 0.3 mm. in
length.
Anteriorly the outer circular muscle is ahout 0.07 mm. in thickness
(PI. 2, Fig 3-CM).
fhe anterior port of the proventriculus is traversed longi­
tudinally hy twelve flattened chitinous teeth Q.2ram. in length and closely set
with small spines (PI. 2, Figs. 1,2,4-Lf).
Beneath the chitinous intima of the
teeth is a continuous layer of small epithelial cell3 (Pi. 2, Fig. 3-BP), and
heneath this, in the cavities of the teeth are strands of longitudinal muscle
(PI. 2 Fig. 3-LM).
In the posterior third of the proventriculus the twelve longitudinal
teeth project into the lumen in the form of rounded ridges (PI. 2, Fig, 4-Lf).
In cross-section (PI. 2, Fig 4) six of these ridges are seen tohe short and
stout, while the six which alternate with them are more slender and project
farther into the lumen,
fhe size of the lumen is thereby much reduced in this
region as compared with its size in the anterior part of the proventriculus.
In the cavities of the teeth are strands of longitudinal muscle (Pi. 2, Fig.4LM).
fhe circular muscle is 0.06 mm. in thickness (CM).
Wu (143) desorihes the proventriculus of UBMOURA vallioularia (Nemouridae)
and indicates that its intima is raised into twelve longitudinal folds, mix
large and six small.
Comstock (32) reports that the food of Pleooptera is
prohahly insects and Grau (60) finds that various species of Perlodes and
Perla feed on larvae of Ephemeri da, Ghironomidae and Gamraarus as well as on
algae.
food,
He concludes that the teeth of the proventriculus act as files on the
fhe examination of the contents of the digestive canal of Aoroneuria
ahnormis revealed the presence of chitinous particles of insects— legs, man­
dibles, etc.
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Ueuroptera
Dafour (42) studied a few species of leuroptera but did not give detailed
descriptions of the proventriouli.
In the larva of Myrmeleon formioarium
(Myrmeleonidae) he found no gizzard, hut in the adult he described and fig­
ured eight chitinous columns running lengthwise in the gizzard and narrow­
ing toward their tips.
He describes the Hemerobridae as having no gizzard.
McDunnough (87) studied the digestive tract of Ohrysopa $>erla L. whose larva
sucks the juices of aphids and litiose adult form eats small insects (119).
The proventriculus of the larva is lined with a thin chitinous intima raised
into irregular longitudinal folds and has an oesophageal valve.
The proven­
triculus of the adult has a thick chitinous intima whioh is organized into
eight longitudinal folds which support a thick covering of spines projecting
into the lumen.
The longitudinal and circular muscles are well developed.
A detailed description of the proventriculus of the carnivorous larva
of Corydalis cornutus h.' (Sialidae) is given by Matheson (86).
has eight longitudinal teeth which are heavily ohitinized.
This organ
Matheson holds
the view that the gizzard exercises a crushing and straining action on the
food.
In this order there is a noticeable difference in the complexity of the
proventriculi of the larvae and adults.
Particularly is this to be noted in
Ohrysopa whose larvae suck the juices of small inseots and whose adults de­
vour aphids, scale insects and mites.
Odonata
Dufour (42) in his study of the digestive tracts of insects speakes of
the HItibellulinesw as having no gizzard.
Bis (106,107) studied a large series
of larvae and adult forms and showed that the lining of the gizzard is made up
of four, eight or sixteen longitudinal folds each of which bears chitinous
teeth of various sizes and number.
Higgins (74) reports that the gizzard of
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soraa forms has as many as thirty-two longitudinal folds.
Galvert (29,30) who
studied Odonata of Mexico agrees with Bis on the morphology of the proventri­
ouli of those forms which "both studied.
Pausaek (52) desorihes in a larva of
Odonata, a heavy valve at the posterior end of the proventriculus which regu­
lates the passage of food into the mid-gut.
Meooptera
IXifour (42) shows the proventriculus of Panorpa oommunia as a slightly swollen
sack, whose inner lining is covered with chitinous hairs.
Bphemerida
Dufour (42) reports that the mayflies have no gizzard,
true hy Heedham, Traver and Hsu (91).
this is shown to he
In the larva of Stenonema sp. there is
sin oesophageal valve, hut no specialization of the proventriculus to disting­
uish it from the oesophagus.
The adult mayflies do not feed and the digestive
tract is atrophied.
Thysanoptera
In Thrips vulgatissima, Klocke (80) shows that there is ho specialization of
the proventricular region.
The intima is thin, and the posterior part of the
proventriculus forms an oesophageal valve.
Triohoptera
Hydropsychidae: Macronema zehratum Hagen.
The proventriculus of M. Zebraturn
is 2 mm. long, cylindrical and occupies the mesothoran and raetathora*.
It con­
sists of two regions separated hy a constriction, an anterior region 0.75 mm.
in diameter and a posterior region 0.15 mm. in diameter.
The anterior region
is surrounded hy a heavy layer of circular muscle 0.1 mm. in thickness (PI. 3,
Pig. 3-CM).
Within this there is a layer of epithelial cells of irregular
shape (SP).
The chitinous intima is thrown up into eighteen longitudinal
teeth, 0.8 mm. in length (PI. 3, Pig. 1,2,3-LT).
Their greatest height is at­
tained at a point about three-quarters of their length from the anterior end
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(PI. 2, Fig. 2,4).
At this point they block the lumen of the proventriculus
when drawn together.
(PI. 3, Fig. 4)
In the posterior region of the proventriculus the circular muscle is
thin— 10 m. in thickness (PI. 3, Fig. 5-CM). In this region the eighteen
longitudinal teeth are continued as folds in the intima "beset with straight
chitinous spines which fill the whole lumen (PI. 3, Figs. 1,2,5-LF, OS).
In
the dorsa4 wall of the hind part of this region the intima and epidermis form
a pad which practically fills the lumen and which tapers posteriorly.
covered with a coating of chitinous hairs (PI* 3, Fig, 2,6-P).
It is
Lateral to the
pad, two longitudinal ridges projeot into the lumen (PI. 3, Fig. 1,6-LB).
In
cross-section the posterior part of the provemtrioulus shows the tapered end
of the pad, and the two longitudinal ridges as projections which hang down as
far as the opposite wall of the proventriculus (PI. 3, Fig. 7-P,LB).
Hydropsyohodes analis Bks. studied hy Branch (24) is a semi-carnivorous
fora and has a well-developed proventriculus with thirty-six longitudinal
teeth.
Glasgow (59) found that the oarnivotrus Hydropayohe oolonioa has 34-38
teeth in its proventrioulus.
Lloyd (82) feports that the larvae of Hydro­
psychidae are carnivorous.
Limnophilidae:
She larvae of this family are herbivorous (Lloyd-82).
Branch (24) describes the larva of Limnophilus indivisus Walker as having no
chitinous teeth in its proventriculus, and Buss (111) shows a similar condi­
tion in the larva of Anabolia laevis Zett.
Phryganeidae:
Lloyd (82) and Gatjen (58) report that larvae of this
family feed on dead plant tissue and on filamentous algae, diatoms and leaves
of living plants.
Phryganea interrupta Say, according to Branch (24) has a
large proventriculus with heavy musculature.
Judging from the findings of these investigators of structure of the
proventriculus and feeding habits of frichoptera larvae it is apparent that
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10 -
the chitinous intima of the proventriculus shows its greatest development in
carnivoarous larvae.
Lepidoptera
The first comprehensive study of the digestive tract of Lepidoptera was
undertaken hy Bordas (19) who investigated several genera in ten families:
Nymphalidae: 4 genera
Sphingidae:
2 genera
Arctiidae:
4 genera
Liparidae:
2 genera
Saturniidae:
2 genera
Cymatophoridae:
1 genus
ftotodontidae: 4 genera
Uoctuidae:
2 genera
Tortricidae: 2 genera
Cossidae:
2 genera
These forms are all herbivorous except two wood-eating forms of the family
Cossidae*
Cossus ligaiperdus Fabr. and Zauzera aesouli L.
foims is there a well-developed proventriculus.
valve in some species.
In none of these
There is a short oesophageal
The chitinous intima is thin and may support small
denticles.
Peterson (98) studied the larva of Protoparce Carolina (Sphingidae) and
showed that it has a short oesophageal valve and no heavy development of the
chitinous lining or musculature.
A smilar condition was found hy Henson (68)
in the larva of Vanessa urtioae (Hymphalidae). Gray (61) studied the proven­
triculus of Homaledra aahalella Chambers (Cosmopterydidae) and found that its
walls were thin and that the chitinous intima was slightly less in thickness
than in the oesophagus.
The musculature was found to he poorly developed.
COLEOPT1BA
Cocindelidae:
In Cicindela 6-guttata Fabr. the proventriculus is oval and
about 2 mms. in length.
gut (PI. 4, Fig. 1-MGP).
It is partially surrounded by the cells of the mid­
In cross-section it shows externally a layer of
roughly cuboidal cells of the mid-gut (PI. 4, Fig. 2-MG).
Beneath this is a
layer of circular muscle (CM), three to four strands in thickness.
The inti-
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-li­
ma Is raised into eight longitudinal folds.
Pour of these are very narrow and
in cross-section are finger-like projections into the lumen (PI. 4, Pig. 2-CP).
Alternating with these are four ridges attached to the wall of the proventri­
culus hy very narrow "bases.
In cross-section these ridges are roughly triangu­
lar, one corner of the triangle projecting into the lumen, and the other two
curled inward (CK).
The cavities of these ridges contain a narrow hand of epi­
thelium (EP) and strands of longitudinal muscle(LM).
of the intima is lined with fine chitinous hairs.
fhe whole inner surface
The posterior region of the
proventriculus is circular in cross-section (PI. 4, Pig. 3).
The finger-like
processes of the anterior region are here replaced hy hroad ridges (CP) and
the triangular ridges of the anterior region are replaced hy rounded ridges (CE).
Carahidae:
Chlaenius sp.
and 3 ram. in length.
In this species the proventriculus is fusiform
At the point of its union with the oesophagus the intima
is raised in four longitudinal folds of irregular shape (PI. 5, Pig. 1-LP).
The cavities of these folds are wellsupplied with
fhe circular musculature is thin-.02
mm. (CM).
longitudinal muscles (LM).
Inthe anterior part of
the
proventriculus the longitudinal folds are rounded and much constricted at their
union with the wall of the proventriculus (PI. 4, Pig. 4, PI. 5, Pig. 1-LF).
They are separated hy low longitudinal ridges (PI. 4, Pig. 4, PI. 5, Pig. 2-LH).
Both longitudinal folds and ridges are clothed with fine chitinous hairs (CH).
A heavy layer of circular muscle surrounds the proventriculus in this region
(CM).
At a point ahout two-thirds of the distance from the anterior end of the
proventriculus the longitudinal folds are reduced in size and the longitudinal
ridges are replaced hy four cushionsof closely packed chitinous spines which
project into the lumen (PI. 4, Pig. 4, PI. 5, Pig. 4. CC).
Posteriorly to
these cushions the longitudinal folds are again prominent and the longitudinal
ridges reappear.
In the region of the oesophageal valve the lumen of the pro—
ventriculus is closed hy the longitudinal folds, and the longitudinal ridges
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are allsent.
12 -
(Pi. 5. Pig. 5).
Braohynus sp.: In this species the proventriculus is fusiform and 1.5 mm.
in length.
ridges.
Its intima has four longitudinal folds separated by low longitudinal
These folds and ridges are clothed with fine chitinous hairs as in
Chlaeniua.
It is simpler than that of Ohlaenius
since there are no chitinous
oushiohs, the folds and ridges being continuous throughout its length.
The structure of the proventriculus of Braohynus and Ohlaenius is similar
to that described in several species of Oarabidae by various authors.
Bordas
(18) describes the proventriculi of several species of Carabus, Oalosoma and
Procrustes.
Schluter (116) described the proventriculus of Carabus auratus.
It is similar to that of Ohlaenius in having four cushions of chitinous spints.
The proventriculus of Sphaeroderus nitidicollis Ohev. is similar to that of
Carabus
(114).
Bess (8) who studied Oalosoma sychophanta h. expresses the
opinion that the intima probably functions in grinding the food while the spines
possibly tend to prevent the food from being forced anteriorly.
Whittington
(133) studied the proventriculus of Harpalus pennsylvanicus Dej, and concluded
from the fact thatit possesses heavy layers of longitudinal and circular mus­
cles, that tt is a powerful organ in the process of digestion.
Dytiscidae:
In this family it is to be noted that in. the larvae which are
sucking Insects, the proventriculus is not well developed, while in the carnivor­
ous adults the proventriculus has a complicated and heavy armature.
(39) shows this in his study of a species of Cybister.
Dogener
The intima of the pro­
ventriculus of the larva is raised into two longitudinal folds, while in the
adult there are eight heavily-chitinized teeth of two types, alternating with
lobes bearing chitinous hairs.
In the adult of Dytisous marginalia L. Bungius
(110) shows that the proventriculus has a structure similar to that of Cybister
and he points out that the larva has no gizzard.
In his discussion of the pro­
ventriculi of adult Goleoptera, Bordas (20) includes the Dytiscidae among those
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13 -
which have a well-developed proventriculus.
In six species of five genera he
describes the characteristic arrangement of eight longitudinal teeth, four
small teeth alternating with fowr large teeth.
The same structure is shown hy
Balfour-Browne (3,4) to he present in seventy-seven species of twenty-two
genera of Dytiscidae.
In some of these the large teeth are oval in shape,
while in others they are triangular.
Hydrophilidaet Bordas (20) inoludes the Hydrophilidae among those forms
which have a well-developed proventriculus.
The larvae of these beetles are
carnivorous, feeding on larvae of aquatic insects, crustaceans, tadpoles and
small fish (Wilson-139) which they tear and mash with their strong jaws.
swallow snails whole, first crushing the shell.
They
The proventriculus of Hydro-
phi ins sp. described by Deegener (38) has the intima and epithelial layer
thrown up into irregular folds, outside of which are the longitudinal and cir­
cular muscles.
The proventriculus converges to a narrow oesophageal valve.
The food is drawn in by a sucking action, and the larva has a complicated suck­
ing apparatus in the pharynx.
Staphyllnidaet
(Pavlovsky-96)
Stophylinua bodices Leo.: In this species the proventri­
culus is about 2 mm. in length and cylindrical in shape.
intima is thrown up into eight longitudinal ridges.
Internally the thin
In the anterior portion,
four of these ridges are larger than the four others which alternate with them
(PI. 7, Fig. 2); in the posterior portion, the eight ridges are of equal size
(PI. 7, Fig. 3, PI. 19, Fig. 1)
In the anterior region the crests of the
ridges are beset with small chitinous spines (Fig. 2)j in the posterior region
there are long chitinous hairs vhich arise from the intima of the sides of the
ridges and between them.
The hairs project into the lumen in eight separate
clusters, meeting each other in the centre of the lumen (Fig. 3-CH). Beneath
the intima is a layer of epithelial cells of irregular shape (Fig. 2,3,4-KP),
and in the cavities of the longitudinal ridges are strands of longitudinal
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muscle (Fig. 2,3-LM).
14 -
Surrounding the proventriculus is the layer of circular
muscle which is three or four strands thick except in the region of the oeso­
phageal valve where it is five or six strands in thickness (Fig. 3,4,5-OM).
In the region of the oesophageal valve the lining is raised in four ridges
which hear no chitinous hairs (Fig. 4).
When flattened out the proventricular
lining has the appearance shown in Fig. 1 with the chitinous hairs (CH) ar­
ranged in eight columns.
The proventriculus in this form is similar to that of Oreophilus villosia
G-rav. described hy Talbot (127).
Silphidae:
The proventriculus species of this family is similar to that
of the Staphylinidae.
Von Langerken.
Several species have been studied by Heymons and
In Phosphuga strata L. (69) which feeds on snails, the intima
of the proventriculus raised in eight longitudinal folds which are beset with
numerous chitinous hairs.
The longitudinal musole is separated into eight
bundles one beneath each fold.
circular muscle.
The whole is sourrounded by a thin layer of
The proventriculi of Ablattaria laevigata F. (70) and of
Silpha oarinata L.
(71) i3 similar to that of P. atrata. All three species
feed on snails*
Dacnidae:
Penthe pimelia Fabr. described by Brubakser has a short oeso­
phagus with its lining raised in longitudinal folds.
As it approaches the
mid-gut it increases in diameter while the relative proportion of the differ­
ent layers remains the same.
Brubaker concludes that "it is doubtful that
this larger portion has any fundtion than conduction of food as it is not
large enough to be called a crop, nor heavily chitinized enough to be called
a gizzard.**
Coocinellidaei
While most speoies of this family are carnivorous preda­
tors, there are a few herbivorous forms.
In those forms which have been
studied, there is to be seen a difference in the development of the proventri-
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15 -
cular region of carnivorous and herbivorous forms.
Ipilachna corrupts Ittuls.
feeds on leaves of beans and its proventricular region described by Potts (103)
and Burgess (28) shows no specialization in structure.
The intima is thin and
in the posterior region bears blunt depressions and spines.
Cerotomegilla fuscllabris Muls.
In the carnivorous
landis (81) describes a pear-shaped proventri­
culus twice the width of. the oesophagus.
The intima supports twelve or more
rows of spines, each row having fifteen to twenty-five posteriorly directed
spines which project into the lumen.
The longitudinal musculature is heavy,
particularly at the posterior end, and the oircular musculature is well-devel­
oped.
Elateridae:
Ludius appropinquans. In this form the proventricular
region shows no specialization in structure to distinguish it from the oeso­
phageal region, except the well-developed condition of the oircular muscles
surrounding the oesophageal valve.
It resembles in structure the proventri­
cular region of Asaphesmemnonius H bst.
described by Bigham (9).
The anterior
portion of the proventricular region has a thin lining of chitin thrown up into
irregular longitudinal folds (PI. 8, Pig. 1-LP).
Beneath the chitnous lining
is an irregular layer of epithelial cells, with poorly defined cell walls. (EP).
Surrounding this layer is the layer of circular muscle, two or three strands in
thickness. (CM).
The posterior region of the proventriculus is of smaller dia­
meter than the anterior region (Pig. 2).
It exhibits the same structure as the
anterior region and in addition bears a few long chitinous spines which arise
from the chitinous lining and vhieh are irregular in their arrangement. (CS).
In the region of the oesophageal valve the epithelial layer shows clearly de­
fined cells, and the circular muscle six to seven strands in thickness (Pig.3).
Pig. 4 shows a cross-section of the oesophageal valve.
the two layers of the mid-gut cells (M&i, MG ii).
Toward the outside are
Within these layers is a
single strand of circular muscle(CM), and a single layer of cubical epithelial
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cells (EP).
16 -
fhe chitinous intima is thin*
Lucanidae:
Bordas (20) in his study of the proventriculi of Ooleoptera
includes the Lucanidae with those forms which have no proventriculus.
This
opinion is "borne out by Patterson (94) who described the digestive tract of
Passalus cornutus Fabr. which feeds on rotten wood.
In this insect the pharynx
bears needle-like teeth on its intima but these are not present in the region
of the oesophageal valve.
Soarabeidae:
rophagous.
The valve is continuous with the oesophagus.
The members of this family are mainly phytophagous and cop-
The greater part of the alimentary canal is made up of mid-gut, the
crop is small or wanting, and there is no proventriculus in forms which have
been studied.
In Phyllophaga gracilis Burm, studied by Fletcher (56), the
posterior end of the oesophagus has a chitinous lining raised in irregular
folds.
Swingle (126) shows that in Popillia japonioa hewm.
the fore-gut is
lined with a folded intima, and that the sphincter muscles of the oesophageal
valve are weak and allow regurgitation of food into the fore-gut.
In the dung
beetle Phanaeus vaindex , Mac L. (Becton-6) the oesophagus is a straight, nar­
row, thin-walled tube, lined with a folded intima and projecting into the mid­
gut as an oesophageal valve.
Another dung-beetle, Canthon laevis Dru. studied
by Cooper (33)has a digestive tract consisting mainly of a long mesenteron, and
having a short oesophagus.
Cetonidaet
These beetles according to Bordas (16) feed on pollen, stamens,
petals and leaves of plants, and are without a gizzard or have a much atrophrid
one.
In Cetonia aurata
the gizzard is slightly swollen and its lining is simi­
lar to that of the oesophagus#
Cerambycidae:
Many of the larvae of this family are wood-borers and
Mansour and Mansour-Bek (84, 85) show an interesting relation between the extent
of development of the proventriculus and the type of food consumed.
Xystrooera globosa
The larva of
eats only sapwood which has a high content of starch and
sugar, and it has no proventriculus.
The larva of Macrotoma polmata eats heart
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wood which has a low content of starch and sugar and high content of cellulose.
Its proventriculus is large and has a thick chitinous lining studded with sharp
uniform teeth,
fhe food must he in a finely difided condition to he digested
and Mansour’s examination of the contents of the mid-guts of Xygtrooera and
Macrotoma shows that the food is more finely divided in the latter than in the
former.
Chrysomelidaet
fhe members of this family are herhivoarus and eat the
leaves of many species of plants,
fhose species whose digestive tracts have
been studied show no specialization in the proventricular region, although the
oesophogeal valve may he well developed.
Altica himarginata, Say studied hy
Woods (141) has a crop which has a thin chitinous intima and which is continu­
ous with the oesophageal valve.
It is surrounded hy a thin layer of longitu­
dinal muscle and a heavy layer of circular musole.
jsribed hy Davidson (35) in Criocerus asparagi, Linn.
A similar condition is desIn this insect, the oeso­
phageal valve consists of six folds which project into the lumen of the mid-gut.
Wilson (141) shows that the proventricular region in Chrysochus auratus Fab. is
short, hut gives no details of its structure.
fenebrlonidae:
fhe published work on the digestive tract of insects in
this family shows that the proventricular region is, not well-develdped.
fhe
whole fore-intestine of the fungus-feeding Boletotherus cornutu3, described hy
Auten (<;) is 3 mm. in length.
spines.
Its intima is thin and hears short chitinous
Longitudinal muscles are in scattered groups outside the epithelium
and a layer of circular muscle surrounds the longitudinal muscles.
Fisher (55)
describes no well-defined proventriculus in the death-watch beetle, Xestohium
rufovillosum DeG. He finds that the crop is usually distended with air and that
in no dissections is any food to he found in the alimentary canal.
He concludes
that the beetle does not feed,
Meloidae:
Macrohasis unicolor K hy.
In this form there are no external
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18 -
constrictions marking the boundaries of the proventriculus except where the
alimentary canal widens abruptly behind the proventriculus to form the mid­
gut.
fhe proventriculus widens gradually from its union with the oesophagus
to the mid-gut.
phageal valve,
Its chitinous lining is thin except in the region of the oeso­
fhe folds of the intima in the anterior part of the proventri­
culus are irregular in number and form.
Posteriorly they are sixteen in num­
ber and Show three different shapes in cross-section (PI. 6, Fig. 2).
four larger folds are roughly mushroom-shaped (MB).
folds are four folds which are finger-shaped (FF).
fh®
Halfway between these
Alternating with these two
types of fold are eight smaller folds, irregular in shape (SF).
Within each of
these folds is a layer of small epithelial cells (EP), and in the larger folds
there are strands of longitudinal muscle (1M\.
fhe whole proventriculus is sur­
rounded by a narrow band of circular muscle (CM),
fhe oesophageal valve pro­
jects into the mid-gut in the form of four long projections (Fig. 1,3-CP).
fhese
are heavily chitinized only at their bases and along the lateral edges of the
basal third,
fhey project into the mid-gut for about one-fifth of its length.
At their extremities they are lightly chitinized at the edge,
fhe pxoventri-
culus of M. unioolor is similar in structure to that of Epicauta cinerea marginata Fab, as described by Everly (50).
Curculionidaet
Hypomolyx piceus, De G,
fhe crop of this insect is large,
and when sedtioned was seen to be packed with sections of plant tissue.
intima is armed with eight lohgitudinal rows of chitinous hairs.
Its
In the pro­
ventriculus these rows of chitinous hairs give place to eight double rows of
rectangular chitinous plates which project into the lumen (PI.9, Fig.l-CP,
PI. 19, Fig. 2).
Close examination of these plates shows them to be composed
of closely appressed chitinous hairs (Fig. 2).
fhe successive plates in each
row are very thin, and touch one another so that each row of from 110-120
plates is only about 1 mm. in length.
Between the two columns of plates which
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make up each row there are long chitinous hairs (Fig. 1,2-CH) similar to those
in the crop.
Beneath the intima is a layer of irregular epithelial cells (IP).
Outside the epithelium and attached to the intima is a layer of circular muscle
(CM) six to seven strands thickness.
gonal.
The outline of the cross-section is octa­
In the posterior region of -the proventriculus the intima is raised in
eight rounded longitudinal folds (Fig. 3-LF).
The proventriculus of Oalandra oryzae described hy MuBray and Tiegs (90)
is similar to that of Hypomolyaca, tut is smaller, having about sixty plates in
each row of the proventriculus.
The proventriculus of G. callosa described hy
Wray (142) is similar to that of C. oryzae.
This structure in these three forms
resembles that of the Scolytidae.
Scolytidaet
In this group the proventriculus shows a striking resemblance
to that of the species studied in the family Gurculionidae with which it is al­
lied.
Fefct (53) has shown by use of photographs of cross-sections of the pro-
ventriouli of several genera, that the lining consists of a series of eight
double rectangular plates, together with long chitinous hairs.
structure of these plates are different in different genera.
The details of
Beal (5) traced
the development of the proventriculus from the first to last instance of
Fityogenes hopkinsi Swaine*
In the adult the plates are slender and tapering
toward the base, rather than rectangular.
several strands in thickness.
The oircular musole is heavy and
Fisher (54) studied Scolytus destructor 01.
and notes that its alimentary canal is of the usual bark-beetle type,
Permaptera
This order is treated as a group of the Orthoptera by some authors such
as Bordas (12, 17) who described the proventriouli of several speoies.
It
resembles that of the Orthoptera in having its intima organized into six
longitudinal plates.
In Forfioula auricularia L. each of these plates is
spatuliform and bears small fihitinous projections.
into the mid-gut to form the oesophageal valve.
The six plates project
The proventriouli of habidura
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riparia ia similar to that of F. auricularia.
Orthoptera
Phasmidae; Diapheromera femorata, Say:
In this insect there is no pro­
ventriculus.
The crop is lined with a -thin intima hearing fine teeth (SI. 10,
Pig. 1-CI).
Beneath this is the epithelial layer (SP) made up of small cubical
cells.
layer is surrounded by strandsof circular muscle(CM).
This
In some
specimens, at the posterior end of the crop there is an invagination of the ventral
wall into the lumen which in longitudinal section appears as a thiok projection
(lnv.)
This
is not a constant feature.
The posterior end of the crop narrows
rapidly toward the oesophageal valve (Oes. V) in which region the intima has no
fine teeth.
Dorsally the oesophageal valve is elongated as a flap which extends
into the mid-gut (FL,MG) for a distance of 1.5 mm.
Bordas (17) studied the crop of Phibalosoma pythonius West, and says that
it is entirely without a masticating armature.
In Aoonthoderus spinosus, Gray
and Necroscia erechtheus Westwood he finds that the posterior part of the crop
is lined with small chitinous teeth.
Sinety (118) studied an inidentified spe­
cies of phasmid and noted the same condition.
In Baoillus rossii, Fabr., Cam­
eron (31) found two dorsal projections of the oesophageal valve into the mid­
gut,' the larger one being outside the smaller.
Mantidae:
Mantis religiosa, L.
In this species ihe proventriculus has
the form of a shallow cone, 2 mm. in length, with its broad end opening from
the crop.
In its anterior region there are six areas of narrow longitudinal
ridges (PI. 11, Fig. 1-AB).
In each of these areas the ridges anastomose
posteriorly toward a point midway in the length of the proventriculus where
they end at a short cluster of chitinous hairs (CC).
In cross-section the
ridges are seen to be narrow and forked at their tips owing to the presence of
grooves which run throughout the length of each ridge (Fig. 3-FT).
Alternating
with the areas of anastomosing ridges are six longitudinal teeth which run the
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21 -
length of the anterior half of the proventrioulus (Fig.l,2-LT). At its pos­
terior end each tooth hears small spines.
Behind each of these teeth there is a
rounded fold which runs to the posterior end of the proventrioulus and is densly
olothed with chitinous hairs.
(Fig. 1,4-LF,CH).
Directly behind each tooth the
ohitinous hairs on the folds form a dense cluster, while posterior to this clus­
ter they may break up into three, four or five oolumns (Fig. 1).
Beneath the
longitudinal teeth there is a layer of epithelial cells of irregular shape
(Fig. 2,3,4-EP) and strands of longitudinal muscle (LM).
the whole proventri-
culus is surrounded by a layer of circular muscles six fibres in thickness,
(Fig. 2,4-CM)*
Bordas (17) described the proventrioulus of Mantis religiosa. His des­
cription differes from the above description in two respects.
He does not show
the longitudinal ridges anastomosing posteriorly, but he does show the ohitinous
hairs in the posterior part of the proventricuius not in six folds, but generally
scattered.
Bordas studied the proventriculi of six species of mantids and ditides
them into two groups:
I-those of simple structure, Mantis religiosa, fenodera
australasiae, Leach, Hierodula bioculata, Bura. and Eremiaphila denticolli3,
Lefebr. and II- those of more complex structure Stagmatoptera predioatoria, Stoll
and S. annulata, Stoll,
The latter differ from the former in having spines on
the anatomosing ridges and in having a denticle rather than a cushion of hairs at
the posterior end of each area of ridges.
Blattidae:
The cockroaches have been used more than any other insects for
the study of the physiology of digestion, and consequently the proventriculi of
the species of this group have been thoroughly investigated.
Bordas (13,17)
divided the species into four groups on the basis of the complexity of structure
of the proventrioulus.
The first group includes species of the genera Epilampra
and Panesthia, These have no chitinous teeth or spines.
Included in the second
group are species of Blabera which have large chitinous teeth at the anterior end
of the proventriculi and longitudinal folds posteriorly.
Ouenot's description
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(54) of the proventrioulus of Eotobia livida, Fabr. is similar to that of the
proventriculi of species in this group.
The third group takes in Polyzosteira
limbata, Burm. which has twelve teeth in six rows, the teeth in each row being
separated by transverse furrows and muscular folds.
Group four includes species
of Blatta, Blatella and Perlplaneta. Each of the six rows of teeth has a large
anterior tooth and a smaller posterior tooth or cushion.
The proventrioulus of
various species of Periplaneta has been studied by several authors:
witsch (99,100), Hamme (104), Sanfbrd (113) and Eidmann (46).
Petrunk-
The proventri-
culus of Blatta orientalis was investigated by Wilde (137) and that of Blatella
germanloa by Boss (109).
Aorldiidae: Dufour (42) desoribea the members of this family as having no
gizzard.
When compared with the elaborate development of the proventrioulus in
other Orthoptera, the development of this region is the Acridiidae is slight.
It
does, however, show organization of
its intima into thesix ridges characteristic
of the orthopteron, proventrioulus.
Several species ofthethree families native
to Horth America were studied.
Oedipodinae:
Dissosteira Carolina L.: In -this speoies there is no external
constriction marking the junction between the crop and proventrioulus.
The former
is about 1 mm. in diameter at its posterior end, and its liningis traversed longi­
tudinally by about forty ridges (PI. 12, Fig. 1-LB, PI. 18. Fig. 2) 0.06 mm. in
height.
Within these ridges are a few small epithelial cells (EP) and fibres of
longitudinal muscle (II!).
The outer layer of the proventrioulus is composed of
circular muscle 0.1 ran. in thickness with five to six layers of fibres (CM).
At
the junction of the crop and proventrioulus the longitudinal ridges are larger,
more irregular in shape and fewer in number than in the crop.
circular muscle (CM) is thinner.
(Fig. 2-LB).
The
In the anterior part of the proventrioulus the
intima has twelve longitudinal ridges which converge posteriorly to form six Y—
shaped ridges (Pi. 18, Fig. 2).
Incross-section the anterior part of the
culus shows the twelve ridges to be
irregular in shape,andtending to be
flattened (ffig. 3-LB). A cross-
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proventri-
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section of the posterior part shows the six ridges containing epithelial oells
(Fig. 4-EP) and longitudinal muscle (LM).
In some cases there are fibres running
transversely from one side of the ridge to 'the other (TM).
fietz, (129) in des­
cribing the alimentary canal of this insect includes the posterior section of the
crop as being part of the proventrioulus.
Enooptolophus sordidus, Burm.: The proventrioulus of this form is similar
to that of D. Carolina, differing only in being slightly smaller in diameter.
The proventrioulus of Oedipoda caerulesoens Lin, has been studied by Dufour
(42), Wilde (137) and Bordas (17) all of vhom daow its structure to be similar
to that of Dissostetra. Bordas also shows the same structure in Oedipoda miniata,
Pallas^Psophus stridulus Lini and Pachytylus cinerascens Fabr.
Locustinae:
In the study of the proventrioulus of the genus Melanoplus
three species which eat different foods were examined:
M. femur-rubrum De G.
which feeds on grasses in fields and pastures, M. bivittatus Say which feeds on
coarse sedges and grasses and M. punctulatus Scudd,which feeds on the needles of
White Pine.
In all three foms the proventrioulus is similar in structure to
that of Disaosteira. In M. bivittatus
its diameter is about 2 mm., in M. femur-
rubrum about 0.7 mm., and in M. punctulatus about 1 mm.
The proventricuii of Melanoplus differentialis Thomas studied by Hodge (77)
and of Aoridium peregrinum, Oliv. A. speoiosum Thumb, and Caloptenus italicus
Lin. studied by Bordas (17) as similar to that of Dissosteira.
Tryxalinaet
Chorthippus curtipennis Harris:
The proventrioulus in this
form is analler than that of other forms studied, being 0.5 mm. in diameter.
It is, however, similar in structure to that of Dissosteira. Bordas (17) shows
in a study of eleven species in six genera of Tryxalinae that these species have
proventriculi with structure like that of Ohorthippus.
Pamphaginae:
Pamphagus elephas Stal
studied by Bordas (16,17) has a pro-
ventriculus which differs slightly from that of members of the foregoing families,
in having six EJ-shaped, rather than Y-shaped ridges at its posterior extremity.
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Pyrgomorphinae: Bordas (17) points out tliat the proventrioulus in Poecilooerus sp. and Pyrgomorpha grylloides Serv. is rudimentary hut it is similar in
structure to that in other Acridiidaft.
The study of Acridudae reveals that the proventriouli in members of this
family show very little variation in form throughout the five sub-families.
In
this respect the Acridudae differs greatly from the Blattidae, Mantidae, Gryllidae and Tettigonudae.
Tettigonudae:
Conooephalinae:
Gonocephalus fasoiatus De G.; In this form the proventri-
culus is 1.5 mm. in length, globular, and joined to the crop by a tubular neck.
Internally this neck supports six longitudinal rows of eight to ten cushions
covered with chiiinous hairs (PI. 14, Fig. 4-C).
Each of the first four of these
cushions bears a ohitinous tooth with many small spines (CT).
Between the longi­
tudinal rows there are four narrow bands of short chitinous hears running longi­
tudinally (CH).
of teeth.
mentary.
The gLobular part of the proventrioulus is occupied by six rows
There are nine teeth in each row (T), the last being small and rudi­
Each tooth has a middle section projecting posteriorly (MT) and bear­
ing several ohitinous spines, and two lateral chitinous lobes also projecting
posteriorly (LL).
On each side of a tooth there is a barbated lobe (BL), roughly
oval in outline and heavily chitinizad.
Between the rows of teeth there are
longitudinal chitinous partitions (Cj*). Each of the rows of teeth has a longi­
tudinal flap projecting into the mid-gut to form the oesophageal valve(0es.V.)
The intima of the proventrioulus is shown in PI. 19, Fig. 3.
A cross-section
of hie neck of the proventrioulus shows the lining in six ridges (PI. 14, Fig.l).
Beneath these ridges is a layer of epithelium (IP) and longitudinal muscle#(EM).
A circular muscle (CM) five strands in thickness surrounds the proventrioulus.
A cross-section of a pro ventricular tooth (Fig. 3) shows the median tooth (MT)
and lateral lobes (LL) containing a mass of spithelial cells (EP), connective
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tissue and bundles of longitudinal mufecle (LM).
The posterior part of the
proventrioulus (Fig* 3) shows the irregular outline of the lohes of the oeso­
phageal valve.
C. brevipennis Soudd;
In this species the proventrioulus is similar to
that of C. fasclatus.
The proventriculi of 0. mandibularis Charp., Salomons megacephala de Haan
and Pseudorhynohus minor Bedteribachee, are similar to that of G. fasciatus with
minor differences in shape of teeth, etc. (17)
Copiphorinaes
Neooonocephalus ensiger Harris!
Here the proventrioulus is
somewhat similar to that of Conocephalus, though larger, being 3.5 mm. in length
Including the neck which joins it to the crop.
In the neck there are six longi­
tudinal rows of pointed chitinous teeth (Pi. 13, Fig. 2-GT).
Posterior to these
teeth, and in the region of the junction of the neck and globular part of the
proventrioulus, there is a series of five cushions covered with chitinous hairs
(C).
Between the rows of chitinous teeth there are four columns of chitinous
hairs as in Oonocephalus (CH).
In the globular part of the proventrioulus there
are twelve teeth (T) in each of the six rows, the last two or three being rudi­
mentary.
The middle section of each tooth (MT) has short spines, and the lat­
eral lobes (LL) are long.
The barbated lobes (3L) are roughly rectangular in
shape and each bears a chitinous tooth.
adjacent rows of teeth.
A chitinous partition (GP) separates
The lobes of the oesophageal valve (Pea. V.) are
roughly triangular in shape and bear chitinous hairs.
In a cross-section
through the middle of the proventrioulus (PI. 13, Pig. 1) six teeth are seen
to fill the lumen.
Each of these shows a sharp median tooth (MT) and a later­
al lobe (LL).
Inside the tooth are spithelial cells (EF) and longitudinal
muscles (LM.).
The whole proventrioulus is surrounded by a layer of ciroular
muscle (CM) ten to twelve strands in Hiickness.
The intima of the proventricu-
lus is shown in (Pi. 18, Fig. 1).
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Phaneropterinae:
26 -
Soudderla ourvioauda:
DeG.
The proventrioulus in this
speoiea is 3 mm. in length, including the neok which is short,
The neck has
six longitudinal rows composed of a few long chitinous teeth (PI* 10, Fig,2-CT)
beneath which are cushions of chitinous hairs (CH).
Posterior to the neok, the
proventrioulus is conical in shape, the broad end being toward the orop,
Its
lining supports six rows of twenty teeth, the last seven or eight of which have
no ohitinous spines but are heavily clothed with hairs.
Each chitinous tooth
has a median portion (MT) which is heavily chitinized * The lateral lobes (LL)
are not chitinized but bear closely set hairs (OS).
The barbated lobes are
likewise beset with hairs (BL)»
The proventriculi of Phaneroptera faloata Scopoli and Acridopeza reticaeelata Guerin are similar to that of Scudderia but have fewer teeth (17).
Ehaphidophorinae:
Ceuthophilus maoulatus Harrist
In this form the pro-
ventriculus is slightly over 2 mm. in length and is cylindrical.
Anteriorly
there are six rows of from six to seven chitinous cushions eaoh (PI. 15-CC).
The first four or five of these bear chitinous teeth with short spines (CT).
In -the main body of ihe proventrioulus there are six rows of teeth with ten
teeth in each row.
hairs (MT).
The median portion of a tooth is made up of a cluster of
The lateral lobes (LL) project backward.
The barbated lobes (BL)
are large and rectangular in outline.
Bordas (17) described the proventriculi in several species of the sub­
families Dectioinae, Pseudophyllinae, Meoopodinae and Epihippigerinae. These
all show conformity to a common plan of arrangement of teeth, but there are
individual differences in families and in genera.
Stenopelmatus fusous Hald.
(Stenopelmatidae) has a proventrioulus vdiich resembles that of the Copiphorinae (Davis -36).
Gryllidae:
Oecanthinae:
Oeoanthus nigricoxnis Walker:
Here the proventrioulus is
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globular in shape, 2 mm. in length aad joined to the crop by a tubular neck*
Internally this neck bears six longitudinal rows of lobes covered with chitinous
hairs (PI. 16, Pig. 4-CL).
In the gLobular part of the proventrioulus these
rows have no chitinous hairs, but have a series of complicated teeth,
tenin
number, the last being reduced in size (Pig. 1-T). A surface view of
oneof
these teeth is shown in PI. 17 Pig. 5.
Pro jecting backward is a flattened me­
dian tooth (MT) bearing five or six pointed projections, or median denticles (MD).
Behind this median tooth are lateral denticles with three or four lobes projecting
laterally (LD).
The posterior part of the tooth is made up of two lateral teeth,
projecting posteriorly and laterally and bearing about fifteen projections (Lf).
Beneath each lateral tooth is a chitinous lobe, or inner barbated lobe (1 BL)
which bears chitinous hairs.
At ihe outer side of each inner barbated lobe there
is a rectangular lobe— the outer barbated lobe (PI. 16, Pig. 4-OBL).
Therows of
teeth are separated by chitinous partitions (PI. 16. Pig. 4-CP). Each ofthe rows
of teeth bears at its posterior end a rounded flap (PI. 16, Pig. 4-0es. V.) which
projects into the mid-gut to form the oesophageal valve.
single tooth are shown in PI. 16, Pigs. 1,2,3.
Cross-sections of a
They show that the tooth is filled
with epithelial cells (EP) and longitudinal muscle (LM), and that the proventri­
oulus is surrounded by six to seven strands of circular muscle (CM).
Oeoanthua niveus De G.;
the proventrioulus of this species is similar to
that of 0. nigricornis.
Gryllinae:
Gryllus pennsylvanious Burm.s
The structure of the proventri­
oulus of 1his species has been worked out by Wilde (137), Bordas (17) and du Porte
(43).
It differs from the proventrioulus of Oecanthus in several respects:
ll) There are eleven teeth in each row.
the median denticles are short (MD)*
lobeless.
(4)
(3)
(2)
In each tooth (PI. 17, Pig, f)
The lateral denticles
(LD) are
The lateral teeth (LT) bear few projections.
Gryllus domesticus Latr.: In this form the proventrioulus is similar to
|h»t,of G. pennsylvanious.
The parts of the teeth are identical in shape and
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size.
28 -
The proventrioulus is, however, more robust in. G. domestious than in
.
a^bpermsylvanious and the chitinous intima is thicker.
Neraobius fosciatus De Gs
The proventrioulus of this species is smaller
than that of Gryllus, and the teeth in the two genera differ.
In contrast to
Gryllus, Nemobidos has fewer median denticles {PI. 17,
Pig. 3-MD) and
broader
lateral denticles (LD).
have 2-3 lobes
coverthe
inner barbated lobes.
The lateral teeth {LTO which
The proventrioulus of S', sylvestris Fabr. desoribedby
Bordas (17) is similar to that of S. faaciatus.
Gryllotaiplnaet
Gryllotalpa sp«; This genus has the largest proventrioulus of any species
studied.
The rows of teeth are 2-2.5 mm. in length, and each has fourteen teeth
(PI. 1®, Pig. l).
In each tooth (PI. 17, Pig. 2) there are ten to eleven median
denticles (MD), broad, truncated lateral dentioles (LD) and rounded unlobed lat­
eral teeth (LT).
(1BL).
The inner barbated lobes are well supplied with ohitinous hairs
The outer barbated lobes are rectangular (PL. 17, Pig. 1-OBL). At the
anterior end of each row of teeth there is a chitinous cushion (PI. 17, Pig. 1-CC)
covered with hairs*
Myrmecophilidaet
Schimmer (115) studied the proventriculi of the minute
crickets M. acervorum, M. oohgaoea and M. nebrascensis.
They are similar in
structure to those of larger oriel©ts but have only fowr to six teeth in each
row.
The foregoing study of the proventrioulus of Gryllidae shows that it is a
well-defined organ in the digestive tract, and is lined with a heavy armature.
Hymenoptera
Tenthredinidae; The larvae of members of this family resemble those of
the Lepidoptera in being continuous feeders and in having no specialization of
the proventricular region.
in its lining.
In Lophyrus sp. the "kaumagen" has a few short spines
(112).
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29
-
Larvae of Heodi prion lecontei, Fitoh., Pikonema aisakensis, Boh., Dip­
rion polytomum Htg,, Arge pectoralis, Leach and Cephaloia sp. were examined.
In all cases the fb re-gut consists of a short narrow oesophagus which joins the
mid-gut in -the mesothoracic segment of the "body.
Internally there is a slight
prolongation of the fore-gut into the mid-gut, and at this point there is a
sphincter muscle,
Formicidae: Dufour (42) describes without detail the gizzard of Formica
pubfescens and F. rufa and regards it as being a triturating organ.
Emery (48)
in studying the proventriculi of ants concerned himself mainly with members of
the families Camponotidae, Dolichoderidae, and Gryptoceridae*
He believed that
the proventrioulus was a pumping organ and shows diagrams to support this con­
clusion.
Mukerji (89) showed that in the ant Dorylus orientalis Westw. the
proventidulus is wanting in -the male and is well developed in the worker which
feeds on earthworms.
inous.
It has a narrow bore and its longitudinal folds are chit­
Forbes (5?) describes in detail the proventrioulus of Oamponotus Lercu-
leanus pennsylvanious de Gag.
In this form -the proventrioulus is well developed
and has a lining of complicated ohitinous teeth,
Vespidae:
Dufour (42) describes with little detail the proventrioulus of
Faspa crabre L. Green (62) shows the proventrioulus of Vespa vulgaris to be an
elongate tube with projections forward and backward into the fore-gut and mid­
gut.
Weil (131) describes a similar structure in Vespa orabro. The posterior
extension of the proventrioulus into the mid-gut is three times as long as the
proventrioulus proper,
Apidaes
The proventrioulus of the honeybee^
Apis mellifica L. has been
studied by several investigators who agree on its structure but not on its
function.
Pavlovsky and Zarin (97) show that its forward projection into the
crop is capable of opening and closing by means of the circular and longitud­
inal musculature,
Whitcomb and Wilson (132) studied the ingestion of pollen,
and have shown that the anterior projection reaches forward into the crop,
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20 -
opens its lumen and draws 1he pollen into the proventrioulus.
In passing
through, the proventrioulus, the grains (are not oraoked as the author states
hut enter the mid-gut whole.
Snodgrass (121) regards the proventrioulus as
being an organ for transmitting food from crop to ventriculus.
He says that
the"four lips spasmodically open with a quivering motion and then roll together
tightly and sink into the mouth."
The proventrioulus of the larva is shown by
fflertat (93) and Svenius (49) to have no specialization.
Other Hymeaoptera studied by Weil (131):
Anthidium manioatum and Heriades
fuliginosus (Megachilidae), Philanthus triangulum (Sphecidae) all have the char­
acteristic tubular proventrioulus with anterior and posterior projections into
the crop and ventriculus#
Bombidae: The digestive tract of the bumblebee Bremus pennsylvanious has
been studied by Swingle (125).
Its proventrioulus is similar in structure to
that of Apis mellifioa.
Weil (131) who studied the digestive tracts of several Hymenoptera says
that, "although the forms studied have many different habits, some social, soli­
tary, parasitic, their digestive canals conform closely to a common structure."
This conformity of structure is to be noted in the proventrioulus of the forms
studied by -the authors mentioned above, and is departed from strongly only in
the ants.
All adult forms studied, including the ants, show a proventrioulus whose
chitinous lining is organized into four plates running longitudinally.
The
anterior end of each of these plates is clothed with small chitinous hairs which
are pointed posteriorly.
All forms show an anterior and posterior projection of
the proventrioulus into the crop and mid-gut.
Weil describes the anterior pro­
jection as a "ventilkopf", the posterior projection as a "ventilschlauch" and
the median part or proventrioulus proper as a "ventilfeals".
In the bees the
ventilkppf is a "honey-stopper" (Whitcomb and Wilson-132) which carries the
food from the crop to the mid-gut.
The ventilschlauch is short in some forms
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31 -
e.g. Heriades fuliginosus (131) and reaches its maximum length in -foe Vespidaft
e.g. Vespa crabro (131).
the mid-gut.
It serves as
a funnel for the food as it passes into
Pavlovsky and Zarin (97) describe it as a structure for preventing
the return of food from the mid-gut into the crop.
In the ants tfctedoes not show
anterior and posterior projections, but is joined to the crop and to the mid-gut
by tubular structures (89).
CORRELATION OF THE STRUCTURE OF THE
PBOVBNTRICULUS WITH FEEDING HABITS
1.
Literature:
Some authors have attempted to correlate the structure of the proventrioulus
with thetype of food eaten by insects, while others have attempted to correlate
it with the manner of feeding.
Bis (106) maintains that the proventrioulus is
not well developed in insects which feed on liquids, while it is well developed
in those which chew solid foods.
Plateau (101), however, points out that many
insects which eat solid foods have no proventrioulus.
In correlating the struc­
ture of the proventrioulus with feeding habits Bis says that insects which have
well developed proventriculi devour their food and then cease feeding, when the
proventrioulus re-chews the food.
Such insects may be called "intermittent
feeders" and are usually active forms.
In inactive insects which feed continu­
ously (e.g. lepidopterous larvae) Bis points out that there is not a well devel­
oped proventrioulus,
2.
Discussion:
It is well known that the proventriculi of insects show a wide variation in
the degree of their development.
This is evident from a study of the foregoing
descriptions of the proventriculi in insects of several orders.
A well-devel­
oped proventrioulus is a definite organ in the digestive tract, being separated
from the crop by a constriction or joined to it by a narrow neck.
Its chit-
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iaous intima ia thick and may consist of two layers.
chitinous teeth and spines.
The intima hears heavy
The longi tudinal musculature consists of several
strands running longitudinally within the cavities of the ridges.
The circular
musculature is heavy and is from five (e.g. Plecoptera) to twelve (e.g. Tettigoniidae) fibres in thickness.
A poorly developed proventrioulus cannot he dis­
tinguished from the crop or oesophagus with which it is continuous.
nous intima is thin and may he raised into long£ tudinal folds.
The chiti­
Longitudinal
muscle may he absent (e.g. Thysanura). When present it consists of a few strands
The circular musculature is thin, being two to four fibres in thickness*
A notable fact is that the majority of insects which have a well-developed
proventrioulus also have a large crop.
the case of herbivorous forms.
Particularly is this to he noticed in
Most of the insects with a poorly developed
proventrioulus have a small crop or none at all.
In the latter case the pro-
ventriculus is continuous with the oesophagus*
Following is a list showing those insects idiich I have studied, in which
the proventrioulus is well developed and those in which it is poorly developed:
A.
Well-developed
B.
Poorly Developed
Plecoptera (larvae
Thysanura
Trichop&era (larvae)
Coleoptera
Coleoptera
Cicindelidae
Elateridae
Orthoptera:
Carabidae
Phasinidae
Meloidae
Acridudae
Staphylinidae
Gurculionidae
Hymenoptera
Tenthredinidae (larvae)
Orthoptera
Gryllidae
Mant idae
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33 -
Tett igoniidae
To supplement these is the following list oompiled from the descriptions
of digestive tracts found in the literature:
Neuroptera:
Ghryaopidae (adults)
Collemhola
Feuroptera
Sialidae (larvae)
Myrmeleonidae (larvae)
Myrmeleonidae (adults)
Chrysopidae (larvae)
Sialidae (adults)
Odonata
Mecoptera
Trichoptera (adults)
Coleoptera
Iphemerida
Dytiscidae (adults)
Thysanoptera
Hydrophilidae
Lepidoptera
Silphidae
Coleoptera
Ceramhycidae (Macrotoma sp.)
Dytiscidae (larvae)
Coccinellidae (carnivorous)
Tenebrionidae
Soolytidae
Ceramhycidae (Xystrocera sp.)
Dermaptera
Chrysomeli dae
Orthoptera:
Scarabeidae
Blattidae
Hymenoptera (adults):
Coccinellidae (herbivorous)
Dacnidae
Tespidae
Cetojaidae
Apidae
Lucanidae
Bombidae
Hymenoptera (larvae)
Group A. is made up oB insects which feed on solid foods whether animal
or vegetable#
This would appear to support Bis* contention that a well developed
proventrioulus is to be found in insects which feed on solid foods.
A study of
Group B., however, shows that there are several classes of insects with poorly
developed proventriculi which feed on solid foods:
Thysanura, Phasmidae, Acrid-
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34 -
iidae, Tenthredinidae (larvae), (hepidoptera (larvae), Oerambycidae (Xystrocera),
Chrysomelidae, Coccinellidae.
It is therefore apparent that feeders on solid
foods do not necessarily have well-developed proventriculi.
As noted previously, Sis says that insects which suck up liquids have no
proventriculi.
fhis was shown to be true in the case of Diptera and Hemiptera
(see introduction). This^is also true in the case of mandibulate insects whose
mouth parts are modified for sacking:
Chrysopidae and Myrmeleonidae (larvae),
Sialidae (adults), Dytiscidae (larvae).
In the Chrysopidae and Myrmeleonidae
the larvae have no proventriculi while their carnivorous adults have.
Sialidae the situation is reversed:
In the
The carnivorous larvae have well-developed
proventriculi while the sucking adults have not.
In Dytisoidae, the sucking
larvae have a poorly developed proventrioulus, and the carnivorous adults have
a well developed one,
A study of the two groups A and B shows that predacious carnivores are
confined to Group A.: Plecoptera, Cicindelidae, Cara'bidae, Gryllidae, Mantidae,
Shrysopidae, Odonata, Mecoptera, Dytiscidae, Hydrophilidae, Blatkidae, etc.
is^these forms that the proventriculus shows its greatest development.
It
In these
there are heavy chitinous teeth in the intima and -there is a heavy musculature,
both longitudinal and circular.
lone of the insects with poorly developed
proventriculi are carnivorous predators.
The fore-going study, therefore, indicates that the nature of the food
eaten can be correlated with the degree of development of the proventriculus in
some cases:
(1)
the proventriculus is well developed in predaceous carnivores
and (2) it is poorly developed in mandibulate sucking insects.
But this does
not provide a basis for -the separation of the two groups A. and B.
If we follow His* classification based on feeding habits, we see that most
of the members of group A. are intermittent feeders on solid foods, while in
Group B, most forms are continuous feeders (except the sucking forms).
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This
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35 -
provides a basis fb r -die separation of Groups k* and B. as far as the investi­
gation has prooeeded.
There are a few exceptions.
One of these is Macrotoma sp*
(Cerambycidae) whose larva is a continuous feeder on hard wood of trees.
has a well developed proventriculus.
This
Three other exceptions are the Ephemerida,.
Tenebrionidae (Xestobium), Trichoptera (adults).
These forms do not feed and have
no proventriculi.
The members of Groups A. and B. can now be arranged to show the relation of
proventrieular development to feeding habits.
Group A. besides including inter­
mittent feeders as mentioned by Sis, also includes continuous feeders on hard­
wood of trees (Macrotoma sp.).
The intermittent feeders can be classed, ft®
(1) carnivorous predators and (2) herbivores.
separated into three classes:
The members of group B. can be
(a) sucking insects (b) non-feeders and (c) con­
tinuous feeders on solid foods*
A.
Proventriculus well-developed
(a) Intermittent feeders
(1)
B. Proventrioulus poorly developed
(a)
Carnivorous predators:
Suoking insects
Neuroptera
Plecoptera (larvae)
Myrmeleoni dae (1arvae)
Heuroptera:
Chrysopidae (larvae)
Chrysopidae (adults)
Sialidae (adults)
Sialidae (larvae)
Coleoptera
Myrmeleonidae (adults)
Trichoptera
Dytiscidae (larvae)
(b)
Coleoptera:
Odonata
Tenebrionidae
Mecoptera
Bphemerida (adults)
Coleoptera:
Trichoptera (adults)
Cicindelidae
Carabidae
Non-feeders
(c)
Continuous feeders on solid food
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36 -
Staphylinidae
Thysanura
Dytiscidae (adults)
Collerahola
Hydrophilidae
Thysanoptera
Silphidae
Lepidoptera (larvae)
Coccinellidae
Coleoptera*
Slateridae
Orthoptera
(2)
Gryllidae
Ceramhycidae (Xystrocera sp.)
Manti dae
Chrysomelidae
Blattidae
Scaraheidae
Dermaptera
Coccinellidae (herhivorus)
Herhivoress
Pacnidae
Coleoptera
Cetonidae
Meloidae
Lucanidae
Curoulionidae
Orthoptera
Phasmidae
Scolyti dae
Orthoptera
Acridiidae
Tettigoniidae
Hymenoptera
Hymenoptera
Tenthredinidae (larvae)
Vespidae
Apidae
Bomhi dae
(h)
Continuous feeders on hard wood
Ceramhycidae
Macrotoma sp.
Carnivorous predators:
As mentioned previously these insects show the
greatest degree of development of the proventriculus.
The chitinous intima has
heavy teeth and spines and the musculature is strong.
There is a well-developed
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crop.
37 -
Each order shows a characteristic grouping of the teeth and spines as
well as a characteristic number of teeth.
In Plecoptera there are twelve longi­
tudinal plates with small spines (PI. 2).
In Heuroptera there are eight longi­
tudinal folds hearing spines (Pg. 7.)
longitudinal teeth (PI. 3).
hearing teeth (Pg. 7).
The carnivorous Trichoptera have eighteen
In Odonata there are from four to thi±ty-two ridges
Panorpa sp. (Mecoptera) has a proventriculus with a
heavy lining of spines (Pg. 8).
In all Coleoptera the teeth and ridges are in
multiples of four, and the posterior region of the proventriculus hears spines
(PI. 4,5,7).
In the Orthoptera the teeth are arranged in six rows (PI. 11,16,17)
as also in the Dermaptera (Pg. 19).
The armature of teeth and spines and heavy musculature in the proventriculi
of this group suggest strongLy that this organ is capable of crushing food.
Evi­
dence that this is the case in Gryllus domeatious Latr. will he given later.
Herbivores:
In this group the chitinous intima of the proventriculus is
thinner than in the carnivores and, while there are chitinous teeth, these are
usually lightly chitinized and covered with spines.
In the Coleoptera, the intima
shows ridges arranged in the characteristic multiples of 4:
lionidae, Scolytidae-8.
rows.
Meloidae-16, Curcu-
In the Orthoptera (Tettigoniidae) the teeth are in six
In the case of Neooonocephalus ensiger which is at times oarnivorous, the
proventrioulus shows the heavy teeth and musculature typical of carnivores (PI.13).
In Scudderia and Ceuthophilus which are herbivorous, the chitinous intima is thin,
and the teeth are crowned with fine hairs (PI. 10,15).
In these herbivores the
crop shows its greatest development, being a thin-walled sac which is easily distenable.
The Hymenoptera, feeders on pollen, are a specialized group in which
the proventriculus is enlohgated and may have a long neck (Formicidae). The
plates of the proventriculus are four in number.
Feeders on hardwood:
In Macrotoma sp. (Cerambycidae) the proventriculus
is well developed and has a heavy intima.
herbivores in having no crop.
It differs from the carnivores and
Mansour and Mansour-Bek report that the proven-
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38 -
triculus is capable of chewing the wood ingested (84).
Sucking Insects!
is no proventriculus.
In the few mandibulate insects which suck up juices there
In the Myrmeleomidae, Chrysopidae and Sialidae the juice
of the prey is sucked up through a channel between 1iie mandibles and maaillae.
In the larva of Dytisous, it is drawn up through a groove in the mandible.
Thetfe
sucking insects have strong pumping mechanisms in the head.
Non-feeders: In these forms the digestive tract is aborted or filled with
air.
In Ephemerida and Trichoptera the mouth-parts are not functional.
Continuous feeders on solid foods:
leaves of trees, grass, etc.:
This group includes mainly feeders on
Lepietoptera (larvae), Chrysomelidae, Coccinellidae,
Phasmida$ Aoridiidae, Tenthridinidae (larvae).
The only well-developed structure
in the proventriculi of this group is the sphincter of the oesophageal valve. •
Some of these forms have a crop while others have the oesophageal valve at the
end of the oesophagus.
The Thysanura and Collerabola, feeders on deoaying material,
algae, etc. have longitudinal folds in the intima and a weak circular muscle.
(PI. 1) (Pg.
).
This group also includes Xystrocera sp. (Ceramhycidae) and
Oossus sp. (Cossidae) which feed on the wood of trees.
They eat the sap wood of
trees, and as shown by Mansour and Mansour-Bek (84) they do not require a pro­
ventriculus for chewing this food.
The Scarabeidae itiich feed on leaves, dung,
etc. have a short fore-gut with no vestige of specialization in the proventricular region.
The Lucanidae feed on decayed wood of stumps and the proventri­
culus is similar to that of Scarabeidae.
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- ,,,/ -.39 FUNCTION OF THE PHOVBJTRICULUS
Literature:
The function of the proventriculus has heen studied hy a great many
authors, and much literature has been published on this subject.
The early
workers examined the proventriculi of large common insects such as, crickets
and cockroaches, and concluded from a study of structure that this organ was
designed to chew and masticate food.
This conclusion led to the application
of the names "gesier", "kaumagen" and "gizzard** to the proventriculus.
Dufour
(42) describes the proventriculus of Orthoptera as "un veritable moulin", and
Dominique (41) says that it serves for trituration of ingested materials.
The opinion that the proventriculus is a treturating organ was challenged
by later experimental evidence.
Plateau (101, 102) studied many Orthoptera,
Coleoptera and Myriapoda and said that the proventriculus is not ”un organe
triturateur".
He based his contention on the fact that many insects which eat
solid food have no gizzard.
Petrunkewitsch (99, 100) agreed with Plateau
that the proventriculi of cockroaches are merely filtering apparatus, and
Hamme (104) reported that in no case is the proventriculus capable of breaking
up small particles, i.e.- it is not a "kaumagen".
Thiel (128) came to the
same conclusion after study of many species of Coleoptera and reports that the
proventriculus is a "Verschlussapparat", while Heymons and von Lengerijfeen
(69,70,71) describe the proventriculus of Silphidae as "ein Beusenapparat".
Eberli (45) reports that the proventriculus of Gryllotalpa is not to be re­
garded as an organ for comminuting food.
Several recant workers do not accept the statement that the proventri­
oulus is incapable of triturating food.
They base their conclusions on the
comparison of the condition of food before and after passing through the proventriculus and on the movements of contraction of proventricular muscles.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
m
-
Bordas reports that the proventriculus is a chewing organ in Dermaptera (12),
Dytiscidae (20) and in those Orthoptera which have well-developed proventri­
culi (17).
Several authors vho hold this view are Davis (36-Tettigoniidae),
Mansour (45-Cerambycidae), Grau (60-Plecoptera), Bidmann (46-Blattidae), du Porte
(43-Gryllidae), Matheson (86-Sialidae), Bungius (110-Dytiscidae), Sis (106- Gryllidae, Carahidae), Bess (8-CaraMdae).
Another function which has heen ascribed to the proventriculus is the pre­
vention of regurgitation of solid foods which have passed into the raid-gut.
This
opinion has heen expressed by Sinety (118) in his study of Phasmidae and by Bordas
(17),
Pavlovsky and Zarin (97) who worked on the bee, Apis mellifera L. consider
that the cardiac valve at the posterior end of the proventriculus prevents regur­
gitation.
Evidence for 1he passage of digestive juices from the raid-gut through the
proventriculus to the Grop has been given by various authors.
Bamme (104) found
secretions of the mid-gut in the crop of several insects, and Swingle (126) noted
the same phenomenon in the Japanese Beetle.
In some specialized insects the proventriculus has been regarded as having
a pumping action, notably in ants studied by Emery (48).
Function of Proventriculus of Gryllus doraesticus hatr.:
In the study of -the function of a well-developed type of proventriculus,
30 specimens of Gryllus domestious Latr. were used.
It was necessary to have
the digestive tracts empty of food before the experiments were undertaken.
Abbot (1) reports that food may remain in the crop for several days in cock­
roaches, while Snipes and Tauber (12QJshow that the average time of passage of
food through the whole digestive tract is 20.6 hours.;^ien captured the crickets
had been feeding on young seedling plants and refuse.
being without food for 36 hours.
Two were opened after
In one of these the crop was empty; in the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
other the crop was half full of plant tissue.
without food for 60 hours.
Three were opened after being
In all three the crop was empty.
of the remaining 25 crickets was fed one meal-worm.
At this time each
These they ate voraciously.
Mealworms were fed because their body walls are well solerotized and solerotized
parts are not digested.
lected.
The pellets of frass egested after feeding were col­
The crickets were opened in lots of four to six at intervals of 3, 3,
12, 23 and 29 hours after feeding.
Hesults of the experiments are outlined in
Table 1.
It is to be noted in the table that the volume of the crops of insects
opened three hours after feeding shows a wide variation (50-196 cu. mm.).
It is
also evident that the food does not pass through the digestive tract at the same
rate in all specimens.
In cricket 2. no food has passed into the mid-gut, while
in cricket 4. food has passed as far as the rectum where a pellet of frass is
forming.
This irregularity in the rate of passage of food is also to be noted
■in crickets opened at different intervals of time after feeding.
A similar ir­
regularity was found by Snipes and Tauber (120) to exist in the passage of food
through the digestive tracts of cockroaches (9-33 hours).
This irregularity
leads to wide variation of volume noted in the crops of crickets opened at equal
intervals of time after feeding.
The crops of all crickets which ate both soft tissues and solerotized parts
contained a loose mixture of these materials.
In these crickets in which food
was present 3-29 hours after feeding, the mid-gut was found to contain a simi­
lar mixture of solerotized particles and soft tissues.
All the pellets of frass,
including those egested as early as 7 hours after feeding, contained chitin.
This shows that there is no difference in the composition of the mixture of food
in the crop and mid-gut and indicates that there is no efficient straining action,
which would retain the solerotized parts in the crop while permitting soft tissues
to pass into the mid-gut.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
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Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.
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•
42
—
In the crops the mixture of cihitin and soft tissues was loose and unpacked.
In all cases the food in the mid-gut was found to he compressed into a cylindri­
cal column surrounded hy the peritrophic membrane.
(PI. 20. Fig. 2).
Is noted
in the table, the column of food is broken up into sections, there being in some
cases as many as five separate columns 6’f food, two in the mid-gut, and one each
in the ileum, colon and rectum.
This indicates that the valve at the posterior
end of the proventriculus acts as a mechanism for regulating the passage of food
into the mid-gut.
The chitin in the crop was in large particles including legs, mandibles,
etc., of the mealworms.
{PI. 20.
Fig. 1).
In the mid-gut the chitinous par­
ticles were smaller in size than those in the crop.
,The possible explanations
of this reduction in size are (1) that the smaller particles of ohitin are pas­
sed through into the mid-gut before the large particles and (2) that the proven­
triculus crushes the food and breaks it up.
It has been shown that there is no
efficient straining action which would allow small particles to pass through the
proventriculus.
Therefore, it is to be concluded that the proventrioulus crushes
the food and breaks it up.
Deegener (40) describes the proventriculus as a "Zerkleinerungsapparat” and
several authors show that it is capable of strong contraction in those forms
which have a strong circular muscle.
Hobson (75) and ?feager (144) noted that
the crop and gizzard of cockroaches and beetles are capable Of strong contraction;
and Berlese (7) and Du Porte (44) show that this contraction of the circular
muscle in Gryllus causes the teeth to come together, thereby closing the lumen,
Sanford (113) contends that the teeth do not fit together closely enough to form
a filter but crush the food.
In the gizzard of Periplaneta Yeager (144) noted
that the teeth, which appear as black dots externally, converge during contrac­
tion and diverge during relaxation of the circular musole.
He also noted that
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
43 -
while the insect is eating the rate of contraction decreases and that when
ingestion is completed the rate of contraction increases several fold.
The results of the foregoing experiments on Gryllus domesticus lead to
the conclusion that while the chief function of the proventricuius is to crush
the food into a column and to regulate its passage into the mid-gut, it is al­
so capable of breaking the particles of food.
Function of Proventricuius in Acridiidae:
As pointed out in the study of the Acridiidae, the proventrioulus in this
group is poorly developed.
Several specimens of Melanoplus bivittatus were di-
sected and the condition of the food in the digestive tract was noted.
In all
cases, the food in the mid-gut, after its passage through the proventrioulus,
was seen to be in a closely packed column surrounded by the peritrophic mem­
brane.
This condition is similar to that found in the food of Gryllus domesti-
ous. Unlike the food of Gryllus, however, the food of Melanoplus showed no indi­
cation of having been broken up.
Within the peritrophic membrane the blades of
grass lay parallel to one another and in a straight column.
In some cases this
column was broken up into two or three sections, indicating that the sphincter
muscle at the end of the proventricuius was capable of pinching the food and al­
lowing it to pass through at intervals.
These observations show that while the
proventriculus of Acridiidae, like that of Gryllidae, paisks the food into a
column and regulates its passage into the mid-gut, it does not crush or masticate.
TAXOIOMIC YA1ME OF THE PBQVMfSICULUS
The diversity of structure displayed by the proventriculi of insects has led
several authors to base keys, for identification of genera, on the characters of
proventriculi and to draw up schemes to show phylogenetic relations between
groups within several orders.
Emery (48) composed a "Stammbaum" to show the re­
lationships between genera of the families Dolichoderidae and Camponotidae of
ants.
Basing his results on the structure of the proventriculi of larvae and
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
44
-
adult dragon-flies, His (106,107) composed a table to show the relationships
existing among eight families of Odonata. Higgins (74) commenting on Bis’s
work says that "classification into genera based on resemblances in gizzard
structure would agree in most cases with that now in use based on structure
of wings and other external features of the body."
'The work of Felt (53) and Swalne (124) on Scolytidae has led these two
authors to conclude that the proventriculi of insects in this group is of
value in toxonomy; and Swaine says, "1 have found
the proventrioulus of
the greatest interest and much pradtical value; but a wider study is apparent­
ly necessary
before definite conclusions can be drawn’
.' In the ^ytisoidae,
Balfour-Browne (3) arranged a key, based on the structure of the proventriculi,
for the separation of twenty-two genera; and in another paper (4) showed that
division of Dytiscids based on structure of this organ coincided with divisions
based on the structure of the tarsal claws.
The complete digestive tract ofOrthoptera has beenused byBordas
17) in classifying the families of this group.
A.
Acolatasia:
His results are as follows:
no diverticula or appendages at fore­
end of mid-intestine.
B.
Atrophied gizzard
BB. Globular gizzard
AA.
Colotasia:
-----
Phasmidae
- - -- -- -- - - Forficulidae
one or more diverticula at anterior end of
mid-gut.
Bl.
Powerful teeth in gizzard; 8 intestinal append­
ages; many Malpighian tubules in eight bunches
------- ... _ _
(14,15,
Blattidae
B2.
Rudimentary gizzard; 8 caeca - - - Mantidae
B3.
Gizzard missing or atrophied; 6 caeca each with
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
45 -
'anterior and posterior projections Acridiidae
B4:
Voluminous gizzard, teeth disposed in six ranks;
2 larger appai&ages at anteii or end of mid-gut
___
B5:
_ _ _ Locustidae
Large gizzard mith powerful amiature; two intes­
tinal appendages; Malpighian tubules open into
"ureter” - -- - - - Gryllidae.
The study of the proventriculi of Acridiidae, pg. has shown no vari­
ations in the structure of this organ on which a key could he hased, except
perhaps size which is rather variable,
Bordas (17) divided the Blattidae
into four groups, the Mantidae into two, the Phasmidae into two, not indica­
ting the differences between genera.
In the Tettigoniidae and Gryllidae
there are great variations in structure of the proventriculi, particularly
in the individual teeth.
Following is a key for the separation of four genera of Gryllidae. Two
species of Gryllus were studied, and two of Oecanthus, but the species in
each of these genera could not be distinguished by examination of the pro­
ventriculi,
A.
Each row of teeth more than 2 mm. in length; 14 teeth
in a row (PI. 17. Fig. 2); median denticles 10 or. 11
(Fig. 2-MD): lateral denticles broad and truncate (LD);
lateral teeth rounded and without lobes (LT)— - - - Gryllotalpa
AA.
How of teeth less than 2 mm. in length; 8-12 teeth in
a row; median denticles 4-6 (Pi. 17. Figs. 3,4,5-MD);
lateral denticles broad or narrow (LD): lateral teeth
with 2 or more lobes (LT).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
B.
46
-
Median denticles long and sharp (PI. 17 Pig. 5-1'®},
lateral denticles with 4-5 lobes (LD); lateral
teeth with about 15 lateral projections (LT) - - - --- ” “ Oecaathus
BB.
Median denticles short and rounded (Fig. 3,4-MD);
lateral projections (LI1)
0.
Lateral denticles truncate and broad (Fig. 3LD); lateral teeth with 2-3 lobes (LT) and
covering the inner barbated lobes IIBL) - - - - - -Hem obi us
CO.
Lateral denticles with rounded points (Fig. 4LD)} lateral teeth with 8-10 lobes (M1) arid not
covering inner barbated lobes (IBL) - - - - - Gryllus.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
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47 -
SU1MAHY
1.
The proventriculi of mandibulate insects in several orders are
studied, anddetailed descriptions of their structure is given*
2.
The insects studied are divided into two groups:
those with well
developed proventriculi, and those with, poorly developed proventriculi.
The
former group includes (l) insects which are intermittent feeders on solid
foods,
trees.
'both carnivores and herbivoresand (2) feeders on the hard wood of
The latter group includes (1) sudsing insects, (2) insects which do
not feed, and (3) insects which are continuous feeders on solid foods.
The
structure of the proventriculi in the insects of each of these groups is cor­
related with the feeding habits of the insects*
3.
An historical survey cf the literature on the function of the pro-
ventriculus is given.
Experimental work was done on Gryllus domesticus Latr*
and indicates that the proventrioulus crushes the food into a column, regu­
lates its passage into the mid-gut and breaks up the particles of food.
of the
Study
food in the digestive tract of Melanoplus bivittatus shows that this
organ squeezes the food into a column and regulates its passage into the mid­
gut.
4.
A survey of the literature on the use of the proventrioulus in taxo­
nomy is given.
Its use in the classification of Orthoptera is illustrated by
a key for the separation of four genera of the family Gryllidae,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
48 -
MATERIALS A HD METHODS.
Insents collected:
Thysanura:
Machilis sp:
Collected near Ottawa, Ont.
Trichoptera:
Sept. 2, 1939 under rocks in woods.
Macronema zelratum Hagen:
Ottawa, Ont.
Sept. 2, 1939 in shallow water of Ottawa River.
This species was identified by Dr. T. H. Prison, Chief of Nat­
ural History Survey, Urbana, Illinois.
Plecoptera:
Acroneuria abnormis
Newm.:
Ottawa, Ont. Sept. 2, 1939 in shallow water of Ottawa River;
found clinging to lower surfaot of rocks.
Identified by
Dr. Prison.
Coleoptera:
Carflbidae:
Ohlaenius sp.:
Ottawa, Ont.
Sept. 2, 1939 under loose rocks close to water’s
edge on Ottawa River.
Braohynus sp.:
Sept.
Aug. 2, 16, 1.939.
Same habitats as Chlaenius sp.
These
two were identified by Mr. W. J. Brown, Entomologist Department
of Agriculture, Ottawa.
Cicindelidae:
Cioindela 6-guttata
Fab.
London, Ont. on gravel road.
Blateridae:
Ludius appropinguana:
Ottawa, Ont.
Meloidae:
June 28, July 3, 9, 18, 1939
Macrobasis unicolor Kby.
Wrightville, Quebec.
Curculionidae:
June 8, 1939 on blue cohosh
Hypomolyn. piceus
Ottawa, Ont.
de G.
June 27, July 22, 1939
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
Staphylinidaei
49
-
Staphylinus "badipes Lee.
Ottawa, Ont.
June 28, July 3, Sept. 6, 1939. in damp woods "beneath
rocks*
Ortheptera:
Phasmidae:
Diapheromera femorata Say:
Ottawa, Ontario, Sept. 5, 1939
Mantidae: Mantis religiosa L.:
Belleville, Ont.
Aug.
3, 15, 1939.
Collected by C. W. Smith,
Entomologist, Dominion Parasite Laboratory, Belleville, Ont.
Tettigoniidae:
Ueoconocephalus ensiger
Ottawa, Ont.
Harris!
Aug. 23, 29, 1939 collected during evening in fields,
Conocephalus fasciatus De G.s
Ottawa, Ont.
Aug. 24, Sept. 9, 1939 in fields.
Conocephalus brevipennis
London, Ont.
Sept. 4,
Soudderia curvicauda
Ottawa, Ont.
Scudds
1939 in sedges and tall grass in swampy areas.
De G.:
Aug. 4, 1939 collected during evening in fields.
Ceuthophilus maculatus
Harris:
June 29, 30, July 14, 16, 1939 under rocks, stumps, etc. in damp and
swampy woods.
Gryllidae:
Oecanthus nigrioornis
Ottawa, Ont.
Walk er:
Aug. 27, 29, Sept. 2, 3, 1939 on raspberry plants and
in flower heads of Zinnia,
0. niveus
Be G.
London, Ont.
Oct. 1939
Gryllotalpa sp.:
Preserved specimens from Ward’s Supply House, lochester, N.Y.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
Hemobius fasciatus
London, Ont.
De G.:
Sept. 6, 1939. Fields and roadsides.
Gryllus pennaylvanious
EogsdLon, Ont.
G. domgstiOus
50 -
Surm:
Sept. 6, 1939.
Fields.
Latr.s
London, Ont.
Mar. 17, 1939 in greenhouse.
Acridiidae:
Dissosteira Carolina
Ottawa, Ont.
L.:
Aug.
Encoptolophus sordidus
Ottawa, Ont.
22,Sept. 13, 1939.
Burm:
Sept. 9, 1939.
Chorthippus ourtipennis
Ottawa, Ont.
Gravel roadways.
Fields.
Harris:
Sept. 9, 1939.
Fields.
Melanoplus femur-rubrum De Geer.:
Ottawa, Ont.
M. bivittatus
Aug. 22, 1939 in fields.
Say:
Ottawa, Ont.
Aug. 29, Sept. 1, 3, 1939.
Found ovipositing in
gravel of roadsides.
M. punctulatus
Scudd:
Ottawa, Ont.
Sept. 3, 5, 1939.
hark of trees and stumps.
Foundovipositing
inoracks
of
This speoies and M.bivittatus were
identified by Professor 1. M. Walker, Department of Biology,
University of Toronto.
Hymanoptera:
Tenthredinidae:
Ueodiprion leoontei
Fitch, Pikonema alaskensis Boh.,
Diprion polytomum Htg., Arge pectoralis Leach, Cephaloia sp.
supplied
hy Dr. A. 1. A. Brown, Entomologist, Forest Insect Survey, Department of
Agriculture, Ottawa, Ontario
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
- 51 -
Histological (Technique:
Digestive tracts of large, heavily ohitinized insects were removed from
insects under water, with the aid of a binocular microscope, and immediately
placed in Bouin’s finative.
Small, soft insects were carefully opened along
the mid-dorsal line and placed in the fiaative.
The tissues were left in the
fixative for from six hours in the case of the anallest insects to twenty
hours in the case of the largest insects.
They were then cleared of fixative
by repeated changes of 70 % alcohol and stored in 70% alcohol.
After being
sectioned at 10M. with a rotary microtome they were stained with Harris* modi­
fication of Ehrlieh's feaemato^ylin and counterstained with eosin.
After this
process, nuclii were stained blue, muscle and connective tissue red, while
chitin remained yellow or brown in colour.
Whole Mounts:
Whole mounts were prepared to show the chitinous lining of the proventriculus flattened out and divested of connective tissue and muscles.
The di­
gestive canal of the insect was removed from the body and pinned down by means
of pins through the crop and mid-gut, to a layer of wax in a Syracuse watch
glass.
The digestive canal was covered with water.
culus was then cut longitudinally with fine scissors.
aid of a binocular microscope.
pulation with fine needles.
The wall of the proventriThis was done with the
The muscular coat was removed by careful mani­
In some eases it could not be removed in this way
so the proventrioulus was placed for twelve hours in a strong solution of po­
tassium hydroxide which dissolved the muscles.
The chitinous lining, divested of muscles was placed in several changes
of alcohol of increasing concentrations to remove water, and finally in abso­
lute alcohol.
The absolute alcohol was washed out with tole»oi, and the chitin­
ous lining was mounted on a slide in Canada balsam or clerite.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
- 52 «*
Photographs:
Photographs of a few types of proventriculi were made.
The digestive
•anal was pinned down and the pioventriculus opened as described in the sec­
tion on whole mounts.
minuten nadeln.
The pxo ventriculus was spread out by pinning with
The muscules were stained purple by the addition of a small
amount of gentian violet.
The preparation was placed under the binocular
microsoope and the lenses focussed carefully on the lining of the proventriculus.
Photographs were taken on standard roll film, 2§-M x li-M, with a camera
placed on one ocular of the microscope#
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-
53 -
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MUKEHJI, D.
1933.
Westiv.
90.
MUBBAY, F. ¥. &
MEGS, 0. W.
1935.
On the Anatomy of the Ant, Dorylus orientalis
Eoologischer Anzeiger.
Metamorphosis of Calandra oryzae.
Journal of Microscopical Science.
91.
92.
Vol. 105.
Vol. 77.
#307N.S.
i\r;3En-iAM, ibaveh &
HSU.
1935.
Biology of Mayflies.
NEWCOMER, E. J.
1914.
Some notes on Digestion and the cell structure
Comstock Publishing Co.
of the Digestive Epithelium in Inseots.
Entomological Society of America.
93.
Quarterly
GSM'SL, 1.
1930.
Vol. 7.
Metamorphosis in the Honeybee.
Morphology and Physiology.
Annals of the
Vol. 50.
#4.
Journal of
#2.
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- 63 -
94,
PATTEiSSON, M. T.
1937.
Cellular Structure of the Digestive Tract
of the beetle Passalus cornutus Pahricius.
An­
nals of "the Entomological Society of America.
30.
95,
BATTUE, 1. s. &
EVAIS, A. M.
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1929.
Insects, Ticks, Mites and Venomous Animals
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PAVLOVSKY, E. N.
1922.
Part. 1#
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Hydrophilus carahoSdes L.
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97.
Pavlovsky, E. 1.
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66.
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its Ferments in the bee-Apis mellifera L.
Journal of Microscopical Science.
98.
PETEBSQM, A.
1912.
America.
PETBUHKEWITSCH,A.
1899.
Vol. 5.
#3*
Zur Physiologie der Verdauung hei Periploneta
zeiger.
PETHUNKEffIl’SCH,A . 1900.
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Annals of the Entomological Society of
orientalis und Blatta germanica.
100.
Quarterly
Anatomy of the Tomato-worm larva, Protoparce
Carolina.
99.
Vol.
Vol.
22.
Zoologischer An­
#583.
Die Verdauungsorgane von Periploneta orien­
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101.
PLATEAU, P.
1875.
fiecherches sur les Phenomenes de la Digestion
chez les Insectes.
Belgique.
102.
PLATEAU, P.
1878.
Vol. 13#
Memoires de l'Academie Hoyale de
Vol. 41.
Beoherches sur les Phenomenes de la Digestion
et sur la Structure de l'Appareil digestif chez les
Myriapodes de Belgiques, Academie Boyale de Bruxelles
Memoires.
vol. 42.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
103.
POMS, S . P.
1927.
Alimentary Canal of the Mexican Bean Beetle
(Epilachna oorrupta muls.).
Yol. 27.
104.
RAMOS, I.
1911.
Ohio Journal of Science
#3.
Die Bedeutung des Proventrioulus hei Coleof-
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105.
RMDOW, E.
1924.
Zur Morphologie und Physiologie des Darmkanals
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Zoologie.
106.
SIS, P.
1897.
Zeitschrift fur wissensohaftliohe
Band.
122
Untersuchung uher die Gestalt des Kaumagens
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Zoologische
Jahrbueher, Ahtheilung fur Systematik.
107.
BIS, P.
Yol. 38
1897.
Vol. 9
Review of "Untersuchungen uher die Gestalt
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108.
B0BEB.TSON, 0. W.
Journal of the Boyal Microscopical Society.
1897.
1936. Metamorphosis of Drosophila melanogaster, in­
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109.
BOS3, H. H.
Journal of Morphology, Philadelphia.
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1930.
Notes on the Digestive and Beproductive
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L.
Transactions of the Illinois State Academy of
Science.
110.
BUSS, E. A.
1908.
Yol.
22.
Die Postemhryonaie.
Entwicklung des Darm-
kanals hei den Trichopteren (Anaholia laevis Zett)
/»
41
Zoologische Jahrhuchez^ Ahtheilung fur Anatomie.
Yol. 25.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
- 65 -
111.
MJNGIUS, H.
1911.
Der Darmkanal der Imago und Larve von
Dytiscus marginalia L.
sohaftliche Zoologie.
Zeitschrift fur wisssnYol. 98.
*1
112.
SAINT-HI LA.IB1,K.
1951.
Uher vorderdarmhange hei Lophyrus-larven und
ihre Bedeutung.
Oekologie.
113.
SANFORD, 1. S.
1918.
Zeitschrift fur Morphologie und
Yol. 21
Experiments on the Physiology of Digestion
in the Blattidae.
Journal of Experimental Zoology.
Yol. 25.
114.
115.
SCHAEFER, P. E.
SCfflivMER, F.
1931.
Alimentary Canal of Sphaeroderus nitidicollis
Chev.
Ohio Journal of Science.
1909.
Beitrag zu einer Monographie der Grylloiden-
gattung Myrmeoophila Latr.
schaftliche Zoologie.
116.
SCJ1LUTER, G.
1912.
Vol. 31.
#5.
Zeitsohrift fur wissen-
Vol. 93.
Beitrage zur Physiologie und Morphologie des
Verdauungsapparates der Insekten. Zeitsohfift fur
Allgemeine Physiologie,
117.
SCHNEIDER, A.
1887.
Uher den Darm der Arthropoden hesonders der
Insekten.
118.
DE S1NETY, R.
1901.
SMIHI, R. G.
1922.
Zoologischer Anzeiger.
Vol. X.
Secherches sur la Biologie et l'Anatomie des
Phasrnes.
1191
Vol. 13.
La Cellule -tome 19
Biology of the Ghrysopidae.
Cornell Univer­
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120.
SNIPES, B. T. &
TAUBER, 0. D.
1937.
memoir 58.
Time required for food passage through the
alimentary tract of the cock-roach, Periplaneta
americana Annals of the Entomological Sociejsy of
America.
Vol. 50.
#2.
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-
121.
SNODGRASS, R. E.
1926.
66 -
Anatomy and Physiology of the Honeybee.
McGraw-Hill Book Co.
122.
123.
SNODGRASS, B. E.
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Alimentary Oanal.
SOMMER, A.
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Principles of Insect
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Canadian Bark Beetles - Part 11.
Preliminary
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SWINGLE, M. C.
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Alimentary tract of the common bumble-bee,
Ohio Journal of Science.
126.
SWINGLE, M. G.
1930.
Research, Washington.
TALBOT, M.
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Anatomy and Physiology of the Digestive Tract
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127.
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Journal of Agircultural
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#3,
Structure of the Digestive System in Creo-
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#5.
128.
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Vergleichende Untersuchungen an den Vormagen-
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129,
TIETZ, H.
1923.
Anatomy of the Digestive System of the
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Entomological Society of America.
130.
TAN GEHUCHTM.A.
1890.
Vol. 16.
#3.
Recherches histologiques sur l’appareil
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La Cellule.
Vol. 6
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131.
WEIL, Ernst
1935.
Vergl eichend-morpho logi sche Untersuchungen am
Darmkanal einiger Apiden und Vespidea Zeitschrift
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132.
WHITCOMB, ¥. &
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1929.
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Mechanics of Digestion of Pollen hy the Adult
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WHITTINGTON, F.B.
1935.
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134.
WIGGLSSWQ'RTH, V. B •
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Research Bulletin.
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Alimentary canal of Harpalus pennsylvanious
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Principles of Insect Physiology.
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135.
136.
WIGGLBSlfORiH,V.B.
1934.
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WILLIAMS, P.X.
1910.
Untersuchungen uher den Kaumagen der OrthoArchiv fur Naturgeschichte.
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WILSON, C. B.
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Anatomy of the larva of Cecidomyia resini-
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139.
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138.
Methuen & Co., London
Annals of the Entomological Society of
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#1
Life History of the Scavenger Water Beetle Hydrous
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Fisheries, Washington.
140.
WILSON, S. J.
1934.
Bulletin of Bureau of
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Anatomy of Chrysochus auratus.
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141.
WOODS, W. C.
1918.
Journal of
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Alimentary Canal of the larva of Altica
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Say.
Society of America.
Annals of the Entomological
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#3,
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142,
1VKAY, B. L.
1937.' Snbry&logy of Calendra callosa.
the Entomological Society of America.
143.
WU, C. P.
1923.
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YEAGEB, J. F.
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Morphology, Anatomy and Ethology of Femoura
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14*.
Annals of
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#3.
Observations on Crop and Gizzard Movements
in the cockroach, Peiiplaneta fuliginosa (Serv.)
Annals of the Entomological Society of America.
Vol. 24.
#4.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE
'Ihysanura:
1.
Machilis sp.
Fig. 1
Cross-section of prothorax
Fig. 11
Cross-section of oesophagus in prothorax
Fig. Ill
Cross-section of oesophagus in posterior part
of prothorax
Fig. IV
Cross-section of mesothorax
Fig, V
Cross-section of oesophageal valve
CM:
Circular muscle
DK:
Dorsal ridge of oesophagus
EP:
Epithelium
1:
Intima of oesophagus
li:
External intima of oesophageal valve
lii:
Internal intima of oesophageal valve
MG:
Oes:
Oes. V.:
VB:
Cells of mid-gut
Oesophagus
Oesophageal valve
Ventral ridge of oesophagus
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T S
Plecoptera:
Pig. 1
11.
Acroneuria abnormis Fewm.
Chitinous intima of proventriculus and oeso­
phageal valve#
Fig. 11
Longitudinal chitinous tooth, showing its
coating of small spinas.
Fig. Ill
Cross-section of anterior region of the pro­
ventriculus
Fig. IV
Cross-section of posterior region of the pro­
ventriculus
CM: Circular muscle
EP: Epithelium
LM: Longitudinal muscle
LT: Longitudinal tooth of proventriculus
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 2.
Fig.4
F ig .3
^
4
: y4-T .r.'
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Trichoptera:
Fig. 1
111.
Maoronema zehratum
Hagen
Chitinous intima of proventriculus and oeso­
phageal valve
Fig. 11
Lateral view of longitudinal tooth of pro­
ventriculus
Fig. Ill
Cross-section of anterior region of the
proventriculus
Fig. 1Y
Cross-section of the proventriculus at a
point three-quarters of its length from
anterior aid
Fig. Y
Cross-section of posterior region of the
proventriculus
Fig. Y1
Cross-section of anterior region of oeso­
phageal valve
Fig. Yll
Cross-section of posterior region of oeso­
phageal valve
CM: Circular muscle
CS: Chitinous spines
EP:
Epithelium
LF: Longitudinal fold
LR: Longi tudinal ridge of oesophageal valve
LT: Longitudinal tooth
P: Pad of oesophageal valve
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Cicindelidae:
Pig. I
IV
Cioindela 6-guttata
Fah.
Cross-section of proventriculus surrounded
“by cells of mid—gut
Pig. II
Cross-section of proventriculus
Pig. III
Cross-section of posterior end of proventri­
culus
CH: Chitinous hairs
CM: Circular muscle
CP: Chitinous fold of proventriculus
CB: Triangular ridge of proventriculus epithelium
1: Chitinous intima
LM: Longitudinal muscle
MG: Cells of mid-gut
P: Proventriculus
Carahidae:
Pig. IV
Chlaenius sp.
Chitinous lining of proventriculus
CC: Cushion of chitinous hairs
LP: Longitudinal fold
Lfi: Longitudinal ridge
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Cara'bidae:
Pig. 1
V
Chlaenius sp«
Cross-section of proventriculus at its
point of union with oesophagus
Fig. 11
Cross-section of anterior region of proven­
triculus
Fig. Ill
Cross-section of longitudinal fold in anterior
region of proventriculus
Fig. IV
Cross-section of middle region of proventri­
culus
Fig. V
Cross-section of posterior region of proven­
triculus
CH: Chitinous hairs
CM: Circular muscle
EP: Epithelium
1: Chitinous intima
LF: Longitudinal fold
LM: Longitudinal muscle
LB: Longitudinal ridge
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
2
PLATE
5.
^
o Zi
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Meloidae:
Pig, 1
VI .
Macrobasis unioolor
Idbey-
Chitinous lining of proventriculus and oeso­
phageal valve
Pig. 11
Cross-section of posterior region of proven­
triculus
Pig. Ill
Longitudinal section of proventriculus and
oesophageal valve
CM: Circular muscle
CP: Projection of oesophageal valve into mid-gut
EP: Epithelium
PP: ♦’finger-shaped” fold of proventriculus
1: Chitinous intima
LM: Longi tudinal muscle
MF: ’’mushroom-shaped” fold of proventriculus
MG: Cells of mid-gut
Oes.: Oesophagus
Oes. V.: Oesophageal valve
SP: Small irregular folds of proventriculus
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P h A T E
Staphylinidae:
Pig.
I
¥11
Staphylinus hadipes
Lee*
Chitinous lining of proventriculus showing
four of the eight longitudinal ridges
Fig.
II
Cross-section of anterior region of proven­
triculus
Pig.
III
Cross-section of middle region of proventri­
culus
Pig.
IT
Cross-section of oesophagealvalve
OH: Chitinous hairs
C1I: Circular muscle
EP: Epithelium
1: Chitinous intima
III:
Longitudinal muscle
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 7.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Elateridae:
Pig.
I
Vlll
Ludius appropinquans
Cross-section of anterior region of pro­
ven trioulus
Pig.
II
Cross-section of posterior region of pro­
ventriculus
Pig.
III
Cross-section of anterior region of oeso­
phageal valve
Pig.
IV
Cross-section of posterior region of oeso­
phageal valve
CM: Circular muscle
OS: Chitinous spine
EP: Epithelium
It Chitinous intima
LP:
Longitudinal fold
MGit Cells of mid-gut
MGii: Cells of mid-gut surrounding oesophageal valve
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
8.
PLATE
*
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Ouroulionidaes
Fig. 1
IX
Bypomolyx piceua
Be G
Cross-seotion of middle region of proventrioulus
Fig. 11
Cross-seotion of one ofthe longitudinal rows
of plates
Fig. Ill
Cross-seotion of posterior region of proven­
triculus
CHs Chitinous hairs
CM: Circular muso1b
CP: Chitinous plate
EP: Epithelium
1: Chitinous intima
LF: Longitudinal fold
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE
Phasmidae;
Fig. 1
X
Diapheromera femorate
Say
Longitudinal section of crop and oesophageal
valve
Gs
Crop
CM:
Circular musole
EP;
Epithelium
FL:
Flap of oesophageal valve
1:
lnv.t
MG;
Oes. V.i
Chitinous intima
Invagination of crop wall into lumen of crop
Cells of mid-gut
Oesophageal valve
Tettigoniidae:
Fig i 11
Scudderia curvicauda
De G,
One of "the six longi tudinal rows of teeth in
lining of proventriculus
BL;
Barbated lobe
CH;
Chitinous hairs
CP;
Chitinous pastition
CS;
Chitinous spines on lateral lobes
CT;
Chitinous tooth
LL;
Lateral lobe of tooth
MT;
Median partion of tooth
Oes. V.s
Flap of oesophageal valve
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE IO.
Fiq.l.
f ’, i
f"v
Fig.2.
UAX_C
Inv.
-Oes.V.
0.7mm.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
0.4 mm.
PLATE
Mantidae:
XI
Mantis religiosa
L.
Fig.
I
Section of chitinous intima ofproventriculus
Fig.
II
Cross-section of anterior region of proven­
triculus through the longitudinal teeth
Fig.
III
Cross-section of part of the wall of the pro­
ventriculus showing the ohitinous ridges
Fig.
1Y
Cross-section of posterior region of proven­
triculus through the longitudinal folds
AB: Anastomosing ridges
CC: Cluster of ohitinous
hairs
CH: Chitinous hairs
CMi Circular muscle
EP: Epithelium
FT:
Forked tip of anastomosing ridge as seen in
cross-section
1: Chitinous intima
LF: Longitudinal fold
LM: Longitudinal muscle
LT: Longitudinal tooth
H:
Cross-section of anastomosing ridge
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLA TE I I.
Fig.I.
0.7 m m
Q.74mm
0.18mm.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T S
Acridiidaes
Xll
Dissosteira Carolina
L.
Pig. 1
Cross-section of posterior region of crop
Pig. 11
Cross-section of anterior region of proven­
triculus
Fig. Ill
Cross-section of middle region of proven­
triculus
Pig. 1¥
Cross-section of posterior region of pro­
ventriculus
CMs
Circular muscle
PP: Epithelium
LIvI: Lon$ tudinal muscle
LE:
LongLtudinal ridge
TM:
Transverse muscle
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 12.
Fig. 2.
Fig.l
r<f'
.CM
JLR
.'M&r
.LM
^ ' r:
#\
p i
■i
:!
■
0.42 mm.
Fig.3.
.CM
£P
.LM
vie.'
..
,'i
VV,~y
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
\:
j
P L A T E
Tettigoniidae*
Pig*
Xlll
Neoconocephelus ensiger
Harris
Cross-section, of middle region of proventri-
1
oulus
Pig.
Intima of proventriculus showing two of the
11
six rows of teeth
BL
Barhated lohe
C
Cushion of chitinous hairs
CH
Chitinous hairs in a column
CM
Circular muscle
CP
Chitinous partition
CT
Chitinous tooth
EP
Epithelium
LL
Lateral lohe of tooth
LU
LongL tudinal muscle
Id
Median section of tooth
Oes. ¥.
Oesophageal valve
One of the teeth in longitudinal row of teeth
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
»■ *' . '4c.
a
' I.
/
)
\
/A
XI A
) ( ) $ (// m (,>
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L 4 f E
Tettigoniidae:
X IV
Con.ocepb.alus fasciatus
De G.
Cross-section of anterior region of proventri-
Fig.
oulus
Cross-section of one tooth in middle region of
Fig. 11
proventriculus
Fig
Fig
,
111
Cross-section of oesophageal valve
IV
Chitinous intima of proventriculus showing two
of the six longitudinal rows of teeth
BL
C
Barhated lobes
Cushion of chitinous hairs
CH
Longi tudinal band of chitinous hairs
CM
Circular muscle
CP
Chitinous partition
CT
Chitinous tooth
BP
Epithelium
LL
Lateral lobe of tooth
LM
Longitudinal muscle
WS
Middle portion of tooth
Oes. V.
IT;
Oesophageal valve
footh of middle region of proventriculus
Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Tettigoniidae:
XV
Ceuthophilus maculatus
Harris
Complete lining of proventriculus showing the six rows of teeth
BL:
Barhated lohe
CC:
Cushion of chitinous hairs
CP:
Chitinous partition
CTs
Chitinous tooth
LL:
Lateral lohe of tooth
MT:
Median portion of tooth
Oes.:
Flap of oesophageal valve
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
TTTfl
8
wysmmmvm
sm['mY>Y>
ih N N b G R SP'
\
1 MG
a n m M g f e
m
s
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Gryllidae:
FigI 1
XVI
Oecanthus nigricornis
Walker
Cross-section of proventricular tooth through
the median portion of the tooth
Fig. 11
Cross-section of proventricular tooth through
the median dentiales
Fig. Ill
Cross-section of proventricular tooth, show­
ing the lateral teeth
Fig. IV
Chitinous intima of the proventriculus, show­
ing ttfio of the six rows of teeth
CL:
Lohe covered with chitinous hairs
CM: Circular muscle
CP:
Chitinous partition
IP: Epithelium
IBL:
Inner harhated lobe
LD: Lateral denticle
II!: Longitudinal muscle
LT: Lateral tooth
Ml: Median tooth
OBL:
Oes. V.:
Outer barbated lobe
Oesophageal valve
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 16.
Fig.I.
Fig.4.
1
,1\ I?''ocV
10; *^ '’?■
\0,o::-/\
:
0.19 m m
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
-Oes.V
P L A T E
Gryllidae:
Pig.
I
X V II
Gryllotalpa
sp.
Chitinous intiraa of proventrioulus show­
ing one of the six rows of teeth
Pig.
II
Proventricular tooth
Wemobius fasciatus De G.
Pig.
III
Proventrioular tooth
Gryllus domesticus
Fig.
IV
Proventricular tooth
Oecanthus nigricornis
Pig.
Latr.
V
Walker
Proventricular tooth
CC: Cushion of chitinous hairs
CP: Chitinous partition
IBL: Inner barbated lohe
LDs
Lateral denticle
LT: Lateral tooth
I®; Median denticle
MT: Median tooth
GBL: Outer harbated lohe
T; Proventricular tooth
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P LA TE 17.
Fig.2.
t\ /i,
i /
> ,\
0.32 mm.
Fig.3.
aaa1
^
sJ'b
_OBL
O.I8mm.
Fig.4.
0.18mm.
: ^ n 0!.
-L L Vi
Fig.5.
/
V..'
M > 0 /
0.7 mm.
'IA A 1
0.14mm.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T E
Tettigoniidaes
Pig.
1
XV111
Ueoconocephalus ensiger
Harris
Chitinous lining of proventriculus
(Spread out and pinned to a sheet of waa)
Acridiidae:
Pig.
11
Dissosteira Carolina
Chitinous lining of proventriculus show­
ing five of the six Y-shaped ridges
(Spread out and pinned to a sheet of wax)
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 18
Fig. I
Fi g.2
r
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A
Staphylinidae:
Fig,
1
® E
XIX
Staphylinus "badipes
Lee.
Chitinous intima of proventriculus spread
out to show the eight rows of chitinous
hairs
Curculionidae:
Fig.
11
Hypomoloxy piceus
De G.
Chitinous intima of crop and proventriculus
spread out to show the eight rows of chitin­
ous hairs and plates
‘Dettigoniidae:
Fig.
Ill
Conocephalus fasciatus
De G.
Chitinous intima of proventriculus
Shese three photographs show the proventrioulus
spread out and pinned to a sheet of wax.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 19
Fig.I
r
7
1
m
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
P L A T S
Pig.
1
Contents of crop of Gryllus domesticus
Latr.
Pig.
11
XX
showing particles of chitin (X13)
Contents of mid-gut of G. domesticus
showing the food compressed into a
column surrounded hy the peritrophio
membrane.
(XI3)
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
PLATE 2 0
r
Fig I
A
Fig. 2
r
'
I
A
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.
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