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Acta Zoologica (Stockholm) 92: 179–186 (April 2011) doi: 10.1111/j.1463-6395.2010.00451.x

Morphology of the stomach of the tropical house gecko

Hemidactylus mabouia (Squamata: Gekkonidae)
Sirlene Souza Rodrigues Sartori,1 Katiane de Oliveira Pinto Coelho Nogueira,2 Alı́pio dos Santos Rocha3
and Clóvis Andrade Neves3

1
Department of Animal Biology – Federal
University of Viçosa, Viçosa, Minas Gerais
36570 000, Brasil; 2Department of Biologi-
cal Science – Federal University of Ouro
Preto, Ouro Preto, Minas Gerais
35400 000, Brasil; 3Department of General
Biology – Federal University of Viçosa, Viç-
osa, Minas Gerais 36570-000, Brasil
Keywords:
morphology, reptile, stomach
Accepted for publication:
3 February 2010
Abstract
Rodrigues Sartori, S. S., Nogueira, K. O. P. C., Rocha, A. S. and Neves, C. A.
2011. Morphology of the stomach of the tropical house gecko Hemidactylus mab-
ouia (Squamata: Gekkonidae). — Acta Zoologica (Stockholm) 92: 179–186.
Hemidactylus mabouia is a common species in Brazil, which facilitates its use in
research in several areas and allows display it as a benchmark for studies with
reptiles. To study the morphology of the stomach of H. mabouia, we carried out
anatomical, histological and histochemical analysis. The stomach of H. mabouia
is ‘J’ shaped and can be divided into oral fundic (OF), aboral fundic (AF) and
pyloric regions. The surface epithelium is composed of mucosecretory cells
(MC) containing neutral mucins. In the lamina propria of the OF region, are
large ramified tubulo-acinar glands, which become smaller, less ramified and
more tubular towards the AF region, and are simple tubular and short in the
pyloric region. The fundic glands contain differentiated neck and pit. The neck
is composed of MC containing neutral mucins and the pit is made of oxyntopep-
tic cells (OC). The OC of the OF region contained many zymogen granules,
while those of the AF region contained few zymogen granules and many mito-
chondria, which suggests the existence of a gradient of pepsinogen and hydro-
chloric acid secretion. Pyloric glands consisted of MC containing neutral mucins
and both argyrophil and argentaffin endocrine cells.
Sirlene S. Rodrigues Sartori, Department of Animal Biology – Federal Univer-
sity of Viçosa, Viçosa, Minas Gerais 36570 000, Brasil. E-mail: rodrigues.sirlene
@gmail.com

aspects are somewhat rarer. The classical references on com-

Introduction
Hemidactylus mabouia is an exotic gecko species and widely
distributed in Brazil. Morphological studies are essential for
understanding the biology of the species, besides promoting
an additional tool to physiological, pathological and phyloge-
netical analyses. Among the morphologic studies, those that
deal with the structure of the digestive system are especially
important to understand the digestive process of the animal
used for its nourishment.
There are few studies investigating the morphology of the
stomach of reptiles and among these, most referred to a spe-
cific aspect of tissue morphology such as location of mucosub-
stances or ultrastructure of certain cell types (Madrid et al.
1989; Arena et al. 1990; Perez-Tomas et al. 1990). Studies
that analyse the morphology of the stomach in a broader con-
text, involving anatomical, histological and histochemical
parative morphology (Andrew and Hickman 1974; Luppa
1977; George et al. 1998) contain very little information on
the morphology stomach of reptiles.
According to Luppa (1977), the stomach of reptiles is
divided into two distinct regions: the fundic region and the
pyloric region. Some studies have shown differences between
fundic and pyloric glands in relation to the morphology and
cellular composition, and the mucins types secreted (Suga-
numa et al. 1981; Madrid et al. 1989; Perez-Tomas et al.
1990; Ferri and Liquori 1992). Other studies have shown dif-
ferences between glands of the oral fundic (OF) region and
the aboral fundic (AF) region with respect to oxyntopeptic
cells (OC), characterizing the existence of a gradient of secre-
tion of proteolytic enzymes and hydrochloric acid along that
region (Lehman and Smith 1975; Giraud et al. 1979; Ferri
et al. 1999; Liquori et al. 2000).

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Acta Zoologica  2010 The Royal Swedish Academy of Sciences 179

Morphology of the stomach of Hemidactylus mabouia • Sartori et al.
Thus, the objectives of this study were (1) describe the
stomach of the gecko H. mabouia from anatomical, histologi-
cal and histochemical analysis, (2) characterize morphological
and histochemical differences between the fundic and pyloric
glands, and (3) characterize morphological and histochemical
differences between the glands along the fundic region, partic-
ularly between the OC.
Material and methods
Fourteen adult specimens of Hemidactylus mabouia (Moreau
de Jonnès, 1818) (Squamata: Gekkonidae), with snout-vent
length of 50–70 mm, males and females, were collected from
January to December 2007 (IBAMA license no.: 10504-1) in
the urban area of Viçosa, Minas Gerais – Brazil (20º45¢14¢¢ S,
42º52¢55¢¢W). After 2 days of fasting, the animals were eutha-
nized with an overdose of pentobarbital injected intraperitone-
ally. Then, the peritoneal cavity was exposed through a
longitudinal incision in the ventral region. All experiments
were carried out according to the ‘Princı́pios Éticos para o
Uso de Animais de Laboratório’ – Ethical Principles for the
use of Animals in Laboratory (Colégio Brasileiro de Experi-
mentação Animal – COBEA 1991). A Vernier calipers were
used to measure the length and diameter of the stomach of
each animal.
For the gross anatomical analysis of the gastric mucosa, the
stomach of four animals was opened along the greater curva-
ture, observed under a stereoscopic microscope (SZ-40;
Olympus, Wetzlar, Germany) and photographed with a cam-
era (Coolpix 4500; Nikon, Tokyo, Japan) coupled to the
microscope.
For the histological and histochemical analyses, pieces
(15 mm2) of the OF, AF and pyloric regions of the stomach
of six animals were collected. The pieces were washed in a
phosphate buffer solution (pH 7.2), fixed in Carson’s forma-
lin solution (Carson et al. 1973) for 24 h, dehydrated in an
alcohol series and embedded in glycol methacrylate-based
resin (Historesin; Leica, Wetzlar, Germany). Using a man-
ual rotating microtome (RM2045; Leica), histological sec-
tions of 3 lm thickness were obtained and stained with
toluidine blue (TB) 1% in distilled water for histological
description or submitted to the following histochemical meth-
ods: alcian blue (AB) pH 2.5 and AB pH 0.5 for the detection
of acid mucins and sulphated acid mucins respectively (Cook
1996); Periodic Acid-Schiff (PAS) for neutral mucins (Pearse
1968); xylidine Ponceau (XP) for proteins (Mello and Vidal
1980); Polak method for mitochondria (Michalany 1981);
and the techniques of Grimelius and Wilander (1980) and
modified Masson-Fontana (Barbosa et al. 1984) for argyro-
phil and argentaffin endocrine cells, respectively, with sections
from the stomach and duodenum of mice used as positive
control.
Stained sections were photographed with a digital camera
(QColor 3; Olympus) connected to an optical microscope
(BX60; Olympus).
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011)
For analysis of the surface morphology of the gastric
mucosa, fragments of the OF, AF and pyloric regions of the
stomach of four animals were collected. The fragments were
washed in a cacodylate buffer solution 0.1 M (pH 7.4), fixed
in a Karnovsky (1965) solution for 24 h, dehydrated in alco-
hol series, critical point dried, coated with gold and examined
with the aid of a scanning electron microscope (SEM; LEO
VP1430 model, Carla Zeiss AG, Jena, Germany).
To obtain the morphometric data, images with the resolu-
tion of 2048 · 1536 pixels were analysed using the IMAGE-
PRO PLUS 4.5 software system (Media Cybernetcs, Inc, Silver
Springs, USA). From the anatomical images was measured
the width of the folds. From the histological and histochemi-
cal images were measured the length of gastric glands and the
thickness of the muscular tunic. Through images of scanning
electron microscopy were measured the width of the small
folds and average width (average between the largest and
smallest widths) of the surface of mucosecretory cells (MC)
and macrophages.
Results
Gross morphology
The stomach of H. mabouia, comprised an OF, AF and pylo-
ric region. The stomach is ‘J’-shaped and is located in the left
antimere (Fig. 1A). It is 19.2 ± 3.0 mm long and at its widest
point, 4.5 ± 1.3 mm. The fundic lumen is wide, relative to
that of the pyloric region. The cranial extremity of the stom-
ach does not have a sphincter and is a continuity of the
oesophagus, while the caudal extremity is extremely con-
stricted and forms a sphincter, the pylorus. The OF region is
sinuous and projects itself to the left antimere; the AF region
is almost rectilinear; and the pyloric region is curved to the
right, towards the midline.
The gastric mucosa is raised into thick, longitudinal and
sinuous folds with rugous surface. In the OF region (Fig. 1B),
about six folds are found, with the width of 0.67 ± 0.08 mm;
in the transition for the AF region (Fig. 1C), some of these
folds disappear, and only two remain in the pyloric region
(Fig. 1D), where they are narrower (0.23 ± 0.03 mm).
Gastric mucosa surface morphology (scanning electron microscope)
In the gastric mucosa of the stomach, besides the thick folds
already described, there are also numerous small folds
(40.9 ± 5.5 lm), ramified and anastomosed, especially in the
AF region (Fig. 2A,B). The luminal surface and the gastric
foveolas (Fig. 2C,D) are formed by MC with convex and
polygonal shaped apex, whose average width is 4.5 ± 1.0 lm.
The apex of the luminal MC is irregularly shaped because of
microprojections of the membrane; the apex of the foveolar
MC is regular (Fig. 2E). Mucus with amorphous aspect or
forming a net was observed on the gastric mucosa
(Fig. 2B,C). Some large cells were also observed on the
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011)
Fig. 1—Anatomy of the stomach of
Hemidactylus mabouia. —A. Stomach with its
oral fundic (OF), aboral fundic (AF) and
pyloric (P) regions. —B. Internal coating of
the oral fundic region. —C. Internal coating
of the transition from the oral fundic region
(of) to aboral fundic region (af). —D. Internal
coating of the pyloric region. Arrows: folds;
LI: large intestine; SI: small intestine; O:
A D
oesophagus.
surface (12.0 ± 1.5 lm of average width), which may be
macrophages. These cells are irregularly shaped and some-
times appeared to produce thin membrane projections
(Fig. 2F).
Histological morphology
In all regions of the stomach the wall is thick and composed of
four tunics: mucosa, submucosa, smooth muscle and serosa.
A single layer of prismatic MC lined the luminal surface and
the gastric foveolas (Fig. 3A–C). These cells contain small
strongly PAS-positive and AB-negative secretion granules,
densely distributed in the supra-nuclear cytoplasm, enlarging
the cellular apex (Fig. 3D–F).
Underlying the epithelium (EP) of the mucosa is the lam-
ina. A basement membrane was not observed. The connective
tissue which fills the lamina propria is sparse and contains
small blood and lymphatic vessels, immune cells, such as mast
cells and lymphocytes, isolated neurone bodies and gastric
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Sartori et al. • Morphology of the stomach of Hemidactylus mabouia
glands (Fig. 3A–F). Sometimes, mitotic figures were observed
among the epithelial cells lining the glandular ducts (Fig. 3F).
In the OF region (Fig. 3A,D), the gastric glands are rami-
fied simple tubulo-acinar, with a large secretory portion
(121.8 ± 34.7 lm). They contain two main cellular kinds:
MC in the neck and OC in the pit. The MC are pyramidal,
with a triangular nucleus located in the basal portion of the
cell and small secretory granules distributed in the cytoplasm.
Secretory granules stain lightly with TB (Fig. 3A) and are
PAS-positive (Fig. 3D) and AB-negative. The OC are polyhe-
dral, and contain an irregularly shaped nuclei located in vari-
ous positions around the cell. They contain large
orthochromatic (Fig. 3A) and XP-positive granules
(Fig. 3G), which are most likely pepsinogen granules.
In the AF region (Fig. 3B,F), the gastric glands are tubular,
smaller (36.2 ± 10.2 lm) and less ramified than those of the
OF. They also contain the cellular kinds previously men-
tioned, but the OC have filiform structures concentrated in
their basal half, which strongly stain with TB (Fig. 3B) and
181

Morphology of the stomach of Hemidactylus mabouia • Sartori et al.
A B
D E
F (OF) region to aboral fundic (AF) region.
show an intense reaction to the Polak technique for mitochon-
dria (Fig. 3H). The OC of the AF region contain a lower den-
sity of zymogen granules than those from the OF region.
In the pyloric region (Fig. 3C,E), the gastric glands are sim-
ple tubular and short (26.7 ± 6.1 lm). They are relatively
homogeneous and mainly composed of MC, which are pris-
matic, tall and have a basally located nucleus. The secretion
granules are reactive only to PAS and are densely distributed
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011)
Fig. 2—Surface morphology of the stomach
of Hemidactylus mabouia. —A. Gastric
mucosa in the transition from the oral fundic
Arrow: thick fold; arrow-head: small fold.
—B. Gastric mucosa of the aboral fundic
region. Arrow: small fold; arrow-heads:
mucus. —C. Mucosal epithelium of the oral
fundic region. Arrows: mucus. —D. Mucosal
epithelium of the aboral fundic region.
Arrows: mucosecretory cells. —E. Crypt of
the aboral fundic region. Arrow:
mucosecretory cell. —F. Mucosal epithelium
of the pyloric region. Stippled circle:
macrophage on the gastric surface.
in the supra-nuclear cytoplasm, except in some cytoplasmatic
portions, where vacuoles appear without being marked by
PAS. The lumen of the pyloric glands is larger than that of the
other gastric glands and sometimes it is possible to observe
the presence of PAS-positive mucus inside them (Fig. 3E).
Endocrine cells were observed in the mucosal and glandular
epithelia throughout the stomach. These cells generally pos-
sessed a clear cytoplasm, not-stained with TB (Fig. 3A,C).
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011) Sartori et al. • Morphology of the stomach of Hemidactylus mabouia

A B C

D E F

G H I

Fig. 3—Histology and histochemistry of the stomach of Hemidactylus mabouia. —A., B. and C. Mucosa of the oral fundic, aboral fundic and pylo-
ric regions respectively (toluidine blue – TB). Arrows: mitochondria in the oxyntopeptic cells; double-headed arrow: gastric gland; EP: epithelium;
GF: gastric fosseta; MC: mucosecretory cells; OC: oxyntopeptic cell; stippled circle: endocrine cell. —D. Gastric mucosa, oral fundic region (peri-
odic acid-schiff, PAS). Arrow-head: mucosecretory cells; EP: epithelium; GF: gastric fosseta; OC: oxyntopeptic cells. —E. Gastric mucosa, pylo-
ric region (PAS). Arrow-head: mucosecretory cells; asterisk: mucus in the glandular lumen; EP: epithelium; GF: gastric fosseta. —F. Gastric
mucosa, aboral fundic region (AB pH 2.5). Arrow-head: mucosecretory cells; EP: epithelium; stippled circle: mitotic figures. —G. Gastric
mucosa, oral fundic region (XP – longitudinal section). Arrows: oxyntopeptic cell; MM: muscularis mucosae. —H. Gastric mucosa, aboral fundic
region (Polak). Arrows: mitochondria in the oxyntopeptic cells; MC: mucosecretory cell. —I. Gastric mucosa, pyloric region (TB). Arrow-head:
serosa tunic; CM: circular muscle; LM: longitudinal muscle; stippled circle: isolated neurone body. —J. Macrophage on the gastric surface (TB).

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Morphology of the stomach of Hemidactylus mabouia • Sartori et al.
These cells were located more frequently in the glands, espe-
cially in the pyloric region. They are large cells, with varied
forms, and their nucleus varies from round to oval and con-
tained uncondensed chromatin, with a visible nucleolus. Both
‘open type’ and ‘closed type’ endocrine cells were found.
These cells displayed argyrophilia and ⁄ or argentaffinity with
the use of Grimelius and Masson-Fontana techniques respec-
tively (Fig. 4A–D).
The muscularis mucosae (Fig. 3G) is well developed,
mainly in the folds, comprising two layers of smooth muscles,
one circular and the other longitudinal. The submucosa tunic
is composed of a sparse connective tissue, with few cell bodies,
large blood and lymphatic vessels and small nerve ganglia.
The muscular tunic is composed of one thick layer of circu-
lar smooth muscle and one very thin layer of longitudinal
smooth muscle (Fig. 3I). The muscular tunic is especially
thick in the pyloric region (360 ± 48 lm). Small nerve ganglia
or isolated neurone bodies are present between the muscle
layers. The serous tunic (Fig. 3I) is thin and composed of a
sparse connective tissue coated by mesothelium. In the lumen
near the coating EP, large cells with irregular shape were
observed, which are probably macrophages (Fig. 3J).
Discussion
As described by Zug (1993) for other reptiles and amphibians,
the stomach of H. mabouia appears as a tube in the shape of a
‘J’, which is wider in the curved area and soon becomes nar-
rower to form the pyloric sphincter. However, different shapes
have been reported for the stomach of other reptiles, for
example, the green iguana Iguana iguana, a herbivorous
A B
C D
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011)
animal, has a ‘U’-shaped stomach (Smith et al. 2001). The
stomach of anuran amphibians (Duellman and Trueb 1985;
Santana and Menin 1994; Castro 1995), which are mainly
insectivorous and swallow entire prey, is similar morphologi-
cally to that of H. mabouia.
The folds observed in the gastric mucosa of H. mabouia,
especially in the fundic regions, are important for the expan-
sion of the stomach during the reception and storage of food.
Although common in vertebrate stomachs, longitudinal folds
may vary in abundance, complexity and appearance (Perez-
Tomas et al. 1990; Santana and Menin 1994; Gallego-Huido-
bro and Pastor 1996; George et al. 1998; Pereira 2000).
In H. mabouia, MC of both the EP and the gastric glands
secrete neutral mucins, which are important for the lubrica-
tion and protection of the gastric mucosa from the hydrochlo-
ric acid and pepsin in the gastric juices (Ferri et al. 1999).
Neutral mucin secretion predominates in the stomach of
many vertebrates, especially amphibians and reptiles (Suga-
numa et al. 1981; Madrid et al. 1989; Perez-Tomas et al.
1990; Gallego-Huidobro and Pastor 1996; Ferri et al. 1999;
Liquori et al. 2002, 2007). However, in stomach of many
mammals, both neutral and acid mucins are produced in simi-
lar quantities (Sheahan and Jervis 1976; Sato and Spicer
1980).
As for most reptiles (Luppa 1977), the stomach of H. mab-
ouia can be subdivided into two regions: fundic region, which
encompasses the majority of the stomach, and a smaller pylo-
ric region. These regions differ primarily with regards to the
morphology of the gastric glands: larger and more complex
fundic glands, with distinct neck and pit, with MC in the neck
and OC in the pit; and shorter and more simpler pyloric
Fig. 4—Endocrine cells (arrows) in the stom-
ach of Hemidactylus mabouia. —A. Argentaffin
cells in the oral fundic gland (Masson-Fon-
tana). —B. Argentaffin cell in the aboral fun-
dic gland (Masson-Fontana). —C.
Argyrophil cells in the pyloric gland (Grime-
lius). —D. Argentaffin cell in the pyloric
gland (Masson-Fontana).
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Acta Zoologica (Stockholm) 92: 179–186 (April 2011)
glands, basically composed of MC (Andrew and Hickman
1974; Lehman and Smith 1975; Perez-Tomas et al. 1990;
Ferri et al. 1999; Pereira 2000).
Like the lizard Chalcides chalcides (Ferri et al. 1999), the
fundic region of the stomach of H. mabouia can be subdivided
into an oral portion and an aboral portion, which mainly differ
in the characteristics of the OC. OC are present not only in
reptiles but also in other non-mammal vertebrates (Arena
et al. 1990; Ruı́z et al. 1993; Grabowsky et al. 1995; Gallego-
Huidobro and Pastor 1996; George et al. 1998). These cells
correspond to the parietal and chief cells of mammals, in that,
they secrete both hydrochloric acid and pepsinogen (Andrew
and Hickman 1974). Although there is a single cell type which
secretes both digestive products, in H. mabouia is possible that
there may exist a secretion gradient of proteolytic enzymes
and hydrochloric acid along the fundic region of the stomach.
In addition, the OC of the OF region contain a greater num-
ber of zymogen granules and certainly secrete more pepsino-
gen, while those of the AF region contain a higher density of
mitochondria and probably release more hydrochloric acid.
The morphology of the OC also changes from the oral to the
aboral region in the stomach of the freshwater stingray Potam-
otrygon sp. (Grabowsky et al. 1995), in the Eurasian green frog
Bufo viridis (Liquori et al. 2002) and in the lizards Anolis caro-
linensis (Lehman and Smith 1975), Tiliqua scincoides (Giraud
et al. 1979) and Chalcides chalcides (Ferri et al. 1999). Accord-
ing to Liquori et al. (2002), the pepsinogen–hydrochloric acid
gradient probably allows the food to be first surrounded by
pepsinogen in the OF region, then as the pepsinogen is con-
verted to pepsin in the acid environment of the AF region,
proteolytic activity begins. This morphological arrangement
reflects that of parietal and chief cells within the gastric glands
of mammals, where pepsinogen produced by chief cells in the
glandular bottom and is then converted to pepsin by secretion
of hydrochloric acid by the parietal cells in the gastric isthmus
and neck (Ito et al. 1985).
According to Grimelius and Wilander (1980) technique,
stains almost all the endocrine cells in the gastrointestinal
tract, except the cells that produce cholecystokinin and
somatostatin, while the Masson-Fontana method stains only
the enterochromaffin cells of type I (stock of serotonin and P
substance) and type II (stock of serotonin and motilin).
According to Fujita and Kobayashi (1977), the endocrine
cells can be classified according to the apical communication
with the lumen: those whose apex reaches the luminal surface
(open type) and those without continuity with the lumen
(closed type). In H. mabouia, both ‘open type’ and ‘closed
type’ endocrine cells, displayed argyrophilia and ⁄ or argentaf-
finity, were found throughout the stomach. It is believed that
the cells of the ‘closed type’ respond to distension or humoral
stimuli, while the cells of the ‘open type’ detect small altera-
tions in the pH or in the composition of the luminal content
(Fujita and Kobayashi 1977). In the stomach of reptiles, the
argyrophil cells are common in the fundic region and in
the pyloric glands, while the argentaffin cells are restricted to
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Sartori et al. • Morphology of the stomach of Hemidactylus mabouia
the superior region of the fundic glands (Luppa 1977). In H.
mabouia, there were no differences in the location or
frequency of these cells along the stomach.
The large cells observed on the gastric surface of H.
mabouia resemble macrophages that may form part of the ani-
mals’ immune defense against invasive microorganisms which
can penetrate the stomach together with food. The presence
of macrophages has not been reported in the lining of the
stomach of other species, although it is common to find these
cells in the lumen of the pulmonary alveoli of mammals
(Andrew and Hickman 1974).
Like most other vertebrates (Luppa 1977; George et al.
1998), the muscularis mucosae of the stomach of H. mabouia
was well developed, formed by two layers of smooth muscle.
In H. mabouia the muscularis mucosae is very thin in the
oesophagus and becomes more developed from the oesophag-
ogastric transition, where glands appear (personal observa-
tion, Rodrigues Sartori, S.S.). The muscularis mucosae is
responsible for the motility of the gastric mucosa (George
et al. 1998) and certainly promotes the activity of gastric
glands.
In the stomach of H. mabouia, the submucous and muscu-
lar tunics are similar to those of other vertebrates (Luppa
1977; Rodrigues et al. 1988; Gallego-Huidobro and Pastor
1996; George et al. 1998; Pereira 2000), although the longitu-
dinal muscle layer is almost imperceptible in relation to the
circular layer in H. mabouia. The submucous nervous plexus
is known to control secretions and blood flux, while the myen-
teric nervous plexus controls the peristaltic movements.
Acknowledgements
The authors express their thanks to Laboratory of Animal
Histology, to Laboratory of Insects Cytogenetics and to
Microscopy and Microanalysis Center of the Federal Univer-
sity of Viçosa, for technical assistance. The authors also thank
the financial support of Fapemig and CAPES.
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Acta Zoologica  2010 The Royal Swedish Academy of Sciences