AJBB Online

2014

Asian Journal Of Biology and Biotechnology

Histology, histochemical and ultrastructural studies on Intestine of

Indian catfish, Clarias batrachus (Linn 1758)
Sudhir G. Chirde1 and Y. A. Gadhikar1*
Department of Zoology, Govt. Vidarbha Institute of Science

Abstract:
The aim of the present study was to describe the histological, histochemical and ultrastructural
studies of the intestine of Clarias batrachus. Information about morphology, histology and
ultrastructural studies is essential for a deeper understanding of the physiological approach of
intestine. The intestinal morphology is influenced by feeding habits. Clarias batrachus being an
omnivorous fish, dwelling in muddy water, show a typical intestinal morphology. Histologically,
the intestine wall of the investigated species consisted of mucosa, sub-mucosa, muscularis and
serosa. Numerous intestinal folds were observed lined with absorptive columnar epithelium along
with absorptive cells and mucous secreting goblet cells. The goblet cells were numerous in the
distal intestine region of Clarias batrachus as compared to middle and proximal region of
intestine. Histological analysis revealed that the intestine PAS and AB positive goblet cells.
Ultrastructural studies of intestine showed the epithelial lining of absorptive cells with regular
microvilli, tight junction complexes, lysosome, microtubules and mitochondria. This study gives
deeper insight on the morphological functional output of the intestine of Clarias batrachus and
will be the basic of comparison with other teleost fishes.
Keywords: Histology, Histochemical, Ultrastructure, , Intestine, Teleost
and Humanities, Amravati (MS), India.
*Corresponding Author: Y. A. Gadhikar, e-mail id.: yash.gadhikar@rediffmail.com

Introduction:
The fish digestive system shows a marked
diversity of morphology and function (Cao and
Wang, 2009; Khojasteh, et al., 2009). Better
information
of
intestinal
morphology,
histological and ultrastructural study is
essential for a deeper understanding of the
physiology and pathology of fish intestine. The
intestine is an organ involved in important
physiological function, being the primary site of
food digestion and nutrient uptake. Therefore
optimum utilization of dietary nutrients
depends generally on the effectiveness of
these functions (Caballero et al., 2003). The
intestinal morphology of fish can be influenced
by feeding habits, food intake frequency, as
well as by body size and shape (Buddington et
al., 1997; Arellano et al., 2002). Depending on
diet, the fish intestine can vary morphologically
from short straight to long coil and complexly
arranged (Govoni et al., 1986; Reinthal, 1989).
The feeding behaviors of fish are classified

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according to the type of food consumed. In

simplest form, fish are either herbivore if they
eat plant material, carnivores if they consume
animal material or omnivores if they eat a
combination of plant and animal materials
(Evans, 1998).
Clarias batrachus (walking catfish) is a
freshwater air breathing species of catfish
found in Central Asia, also known in English
walking catfish, and in Marathi Mangur, its
Walk is wriggling motion with a snakelike
movement, by using its pectoral fins to keep it
upright. In the wild, the natural diet habit of this
creature is omnivorous (Mills, 1993). Walking
catfish has an elongate body shape. This
catfish has long-based dorsal and anal fins as
well as four pairs of sensory barbells, and the
pectoral fins contain poisonous spines which
are especially stout on the male.
The present study aimed to study the general
histology and histochemical and ultrastructural

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Histology, histochemical and ultrastructural studies on Intestine
of Indian catfish, Clarias batrachus (Linn 1758)

distribution of mucosubstances in the intestine

of walking catfish Clarias batrachus. This study
gives a deeper insight into the morphological
and functional aspect of the intestine of Clarias
batrachus and will be the basis of the
comparison with other teleostean fish.

Material and methods

1 Collection and samples:Walking catfish
Clarias batrachus (Linn, 1758), (Order:
Siluriformes, Family: Clariidae) were used in
the present investigation. The adult walking
catfishes, ranging from 25 to 30 cm in length
and 250 to 300 gm in weight, were collected
alive from the fish market from the Amravati
city (Maharashtra, India). They were killed and
dissected immediately in the laboratory.
2 Histological, histochemical and
ultrastructural studies
A. Histological and histochemical study:
Fresh adult specimens were carefully
dissected mid-ventral incision and small pieces
of intestine removed from the body. Small
pieces of intestine viz. proximal, middle and
distal parts of the intestine were fixed by
immersion in 10% Formalin or Bouins fixative
for light microscope studies. Samples were
routinely processed by dehydration in an
ethanol series and then embedding in paraffin.
Six micrometer thick histological sections were
cut by microtome, prepared according to
standard protocol and then stained with
haematoxylin and eosin (H and E).
Histochemical techniques were applied for the
identification of mucous in the alimentary tract
of the Clarias batrachus. Section were stain
with Periodic acid Schiff and alcian blue (PASAB) to demonstrate the full complement of
tissue proteoglycans and alcian blue (AB) stain
specifically acidic mucins while PAS stain
neutral one. Micrographs were taken with an
Olympus microscope.
B. Ultrastructural study: Small pieces of the
proximal, middle and distal intestine tissues of
Clarias batrachus were cut and fixed into the
2.5% glutaraldehyde as well as 2%
paraformaldehyde in 0.1 M sodium phosphate
buffer saline solution at pH 7.3 for an about 12
0
hours at 4 C. Samples were post-fixed in 1%

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OsO4 for 1 hour at 4 C. The samples were

dehydrated in an ascending grade of acetone,
infiltrated and embedded in araldite CY 212
(TAAB, UK). Thick sections (1 m) were cut
with an ultra-microtome, mounted onto glass
slides, stained with aqueous toluidine blue and
observed under a light microscope for gross
observation of the area and quality of the
tissue fixation. For electron microscope
examination, thin sections of grey-silver colour
interference (70-80 nm) were cut and mounted
onto 300 mesh- copper grids. Sections were
stained with alcoholic uranyl acetate and
alkaline lead citrate, washed gently with
distilled water and observed under a Morgagni
268D transmission electron microscope (Fei
Company, The Netherlands) at an operating
voltage 80 kV. Images were digitally acquired
by using a CCD camera (Megaview III, Fei
Company) attached to the microscope (AIIMS,
New Delhi, India).
Result
A. Histology and Histochemical study:
The intestine of Clarias batrachus divided into
three parts viz; proximal, middle and distal
intestine. The intestine wall layers which are
divided into mucosa, submucosa, muscularis
and serosa are similar to the other vertebrates
(Fig. 1). The mucosal surface of proximal
intestine part had numerous elongated and
deep finger-like folds called villi, lined by a
simple columnar epithelium consisting of
absorptive cells and mucus secreting goblet
cells (Fig. 2). The folds present in middle and
distal part of intestine are fewer and shorter.
There was tremendous increase in number of
goblet cells observed in the present study in
the distal part of intestine showing high
mucous secretion, as compared to proximal
and middle part of intestine i.e. duodenum
(Fig. 3, 4, 5). The goblet cells appeared like a
swollen circular-ovoid shaped which open on
the luminal exterior by a single pore. The free
borders of the columnar cells outline like a
brush border, which continue to the columnar
epithelium and interrupted only by the opening

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Histology, histochemical and ultrastructural studies on Intestine
of Indian catfish, Clarias batrachus (Linn 1758)

of the goblet cells. Many lymphocytes usually

usually were observed to get aggregate at the
bases of the columnar epithelium while some
of them often appear in the free ends of the
cells. The lamina propria consists of loose
connective tissue that penetrates as well as
supports the intestinal villi. The submucosa of
the intestine is formed of a vascularized loose
connective tissue that lies below the lamina
propria without separating line (Fig. 6). The
muscularis of proximal, middle as well as
posterior intestine consists of two layers of
smooth muscle fiber; thick inner circular and
outer longitudinal muscle layer (Fig. 7). Many
nerve plexuses lie between these two muscle
layers. The serosa is an extremely thin layer of
a simple squamous epithelium. These cells are
flattened with small amount of cytoplasm and
compressed oval nuclei. In proximal, middle as
well as the distal intestinal part of Clarias
batrachus, the mucosal epithelium consists of
a series of goblet cells which were stained
strongly with positive AB and PAS (Fig. 8 and
9). This indicated the presence of the acidic as
well as neutral (carboxyl and phosphate
group) glycoprotein.
B. Ultrastructural study: The ultrastructural
studies of intestine of adult Clarias batrachus
(Control) were carried out in the present
investigation. TEM study showed the apical
border of the proximal epithelial lining of
intestine with presence of microvilli with normal
length, and no clear terminal web below the
microvilli (Fig. 10). Fine filaments of the
microvilli extended in to the apical cytoplasm.
Many longitudinal oriented microtubules, small
vesicles and multivesicular bodies are present
in the apical cytoplasm (Fig. 11 and Fig. 12).
Below the microvilli tight junction are present
(Plate Fig. 13). The endocrine cells with large
oval or spherical shape nucleus with prominent
nucleoli were located either at the base or the
center of the cell (Fig. 14 and Fig. 15). Ovoid
mitochondria were located to be scattered
throughout the cytoplasm and basal to the
nucleus. The rough and smooth endoplasmic
reticulum located near the nucleus of cell (Fig.
15). Lamellar structures were observed in the
cytoplasm of columnar epithelial absorptive
cells (Fig. 13). The apical end of middle
intestine bears numerous microvilli, tight

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junction,
and
microtubules
(Fig.
16).
Ultrastructuraly, long columnar epithelial
absorptive cells were observed in intestinal
mucosa of C. batrachus (Fig. 17). The bulky
goblet cells were filled with numerous mucous
droplets of low electron-density between
surface epithelial cells (Fig. 17). The epithelial
mucosa of intestine contained many endocrine
cells. They were characterized by irregular
hyper-chromatic nucleus and electron dense
granules (Fig. 18). The enterocytes were
joined to the apical surface by the tight
junction complexes including the zonula
occludents and desmosoma (Fig. 16 and 19).
The distal intestine also showed apical end
covered with many microvilli, just below the
microvilli tight junction complexes and some
microtubules were presence adjacent to goblet
cell (Fig. 20). Some round, elongated and
irregular type of enteroendocrine cells could
be established in the distal region of the
intestine. These cells show presence of narrow
extension which is known as cytoplasmic
process and it contains longitudinal oriented
microtubules (Fig. 21 and 22). Based on the
transmission electron microscopically data,
some elongated and irregular open type of
enteroendocrine cells could be established in
the proximal, middle as well as in the distal
intestine. Whereas some Closed type cells
are observed in the middle and proximal
intestinal region (Fig. 14).
Cell type I with round morphology having the
smallest secretory granules were located in
proximal, middle and distal parts of the
intestine (Fig. 14). Cell type II, with elongated
and round shape with secretory granules of
normal size were found in the proximal as well
as the middle parts of the intestine region (Fig.
15 and 22). Cell type III with secretory
granules of nearly same size as those of the
Cell type II but with an irregular shape found in
proximal, middle as well as distal intestinal
region (Fig. 19 and 21).
All the cell types seem to be open type that
means they have a secretory granules
containing base and long narrow extension to
the lumen. This dendrite-like cytoplasm
contains longitudinally oriented microtubules.
Slight pinocytotic activity is observed in the
apical part of the endocrine cell, in which the

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Histology, histochemical and ultrastructural studies on Intestine
of Indian catfish, Clarias batrachus (Linn 1758)

Figure 1-9: HE and PAS, AB study of

proximal, middle and distal intestine of
Clarias batrachus.
Fig. 1: Proximal part of intestine of Clarias
batrachus showing: MF (mucosal fold), M
(Mucosa), LP (Lamina propria), SM (Submucosa), CML (Circular muscle layer), LML
(Longitudinal muscle layer), S (Serosa). (HE
X10). Fig. 2: Mucosa of proximal intestine
showing: absorptive cell (arrowhead), goblet
cell (arrow), LP (Lamina propria). (HE X40).
Fig 3: Middle part of intestine Cl. batrachus
showing: M (Mucosa), SM (Sub-mucosa), LP
(Lamina propria), ML (Muscularis), GC (Goblet
cell). (HE X10). Fig. 4: Distal part of intestine
Cl. batrachus showing: M (Mucosa), SM (Submucosa), LP (Lamina propria), MF (Mucosal
fold, white arrow), ML (Muscularis). (HE X10).
Fig. 5: Mucosa of distal intestine showing:
goblet cell (arrow), LP (Lamina propria). (HE
X40). Fig. 6: Basal part of Intestinal fold
showing: GC (Goblet cell), LP (Lamina
propria), SM (Sub-mucosa), Loose connective
tissue (arrow). (HE X40). Fig. 7: Muscularis
layer of intestine showing: CML (Circular
muscle layer), LML (Longitudinal muscle
layer), Loose connective tissue (arrow). (HE
X40). Fig. 8 and Fig. 9: Mucosa of Intestine
showing: PAS and AB positive GC (Goblet
cell), LP (Lamina propria). (PAS and AB X40).

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Histology, histochemical and ultrastructural studies on Intestine
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Figure 10-22: Ultrastructure of Proximal,

middle and distal intestine of Clarias
batrachus.
Fig. 10: Ultrastructure of epithelial lining of
proximal intestine showing: MV (Microvilli).
Scale bar 500nm. Fig. 11: Microtubule
extends towards the microvilli. Scale bar
200nm. Fig. 12: Many longitudinal oriented
microtubules (arrowhead), small vesicles
(white arrowhead), multivesicular bodies
(arrow) and mitochondria in the apical
cytoplasm. Scale bar 500nm. Fig. 13: Apical
end
showing
tight
junction
(arrow),
microtubules
(arrowhead)
and
lamellar
structure (LS). Scale bar 1m. Fig. 14:
Endocrine cell showing: endoplasmic reticulum
(ER), L (Lysosome), Nucleus (N) electron
dense granules (arrow). Scale bar 500nm. Fig.
15: Endocrine cell showing: Nucleus (N),
Mitochondria (MT), Endoplasmic reticulum
(ER), electron dense granules (G). Scale bar
500nm. Fig. 16: Epithelial border of middle
intestine showing: Microvilli (M), Tight junction
(arrow), Microtubules (arrowhead), Enterocyte
cytoplasm (ET CP). Scale bar 2m. Fig. 17:
Apical end of middle intestine showing: Goblet
cell (GC), microvilli (MV), absorptive cell (AC),
nucleus (N), mitochondria (MT). Scale bar
2m. Fig. 18: Endocrine cell show irregular
hyper-chromatic nucleus (N) and electron
dense granules (G). Scale bar 1m. Fig. 19:
Irregular shape endocrine cell showing:
Nucleus (N), mitochondria (MT), lysosome (L)
and vacuoles (V). Scale bar 500nm. Fig. 20:
Apical end of distal intestine showing: microvilli
(MV), tight junction (arrow), microtubule
(arrowhead), goblet cell (GC). Scale bar 2m.
Fig. 21: Endocrine cell with elongated
cytoplasm
showing:
Nucleus
(N),,
Mitochondria (MT), Cytoplasmic process (CP),
Microtubule (arrowhead), Golgi body (GB).
Scale bar 2m. Fig. 22: Endocrine cell from
distal intestine region showing: Nucleus (N),
long cytoplasmic process (CP), electron dense
granules (G), microtubules (arrowhead). Scale
bar. 2m.

Discussion:
A. Histological and histochemical study:
The histological studies of the digestive tract

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Histology, histochemical and ultrastructural studies on Intestine
of Indian catfish, Clarias batrachus (Linn 1758)
across species of fish are more valuable as
the interest in fish culture expands and
therefore more information necessitated with
regard to feeding and nutrition. The intestinal
mucosa acts as selective barrier to nutrients
and also prevents many toxins and pathogens
(Murray et al., 1994). Moreover, it plays a part
in the endocrine function, electrolytic balance
and immune response (Buddington et al.,
1997). The fish species in present study
belongs to the family clariidae, showed
oesophageal wall with a more regular structure
and longitudinal folds, which is related to the
oesophagus of the silverside odontesthes
bonariensis (Diaz et al., 2006) and more
simple than those found in marine teleosts
(Abaurrea-Equisoain
and
Ostos-Garrido,
1996).
The intestine of Clarias batrachus is divided in
to three part; proximal, middle and distal
intestine. Long, numerous mucosal folds were
observed in the proximal intestine increasing
the surface area and enhancing the absorptive
activity (Khanna, 1971), while short in middle
and distal region. The mucosal folds of
intestine lined by a single layer of columnar
cells with numerous mucous secreting goblet
cells were consistent and in agreement with
the results obtained in other species
(Khojasteh et al., 2009; Abdulhadi et al., 2005;
Cataldi et al., 1986; Hernandez et al., 2009).
Columnar epithelial cells of the intestinal
mucosa may have an absorptive function as
reported in other fishes (Sis et al., 1979;
Clarke and Witcomb, 1980; Albrecht et al.,
2001). The pattern of muscularis layers
consisting of thicker inner circular and outer
longitudinal smooth muscle observed in
Clarias batrachus. This was also found similar
to that described for Ambassis sp. (Martin and
Blaber, 1984), for Tilapia (Gargiulo et al.,
1998), for Orthrias angorae (Sucmez and Ulus,
2005) and for Rhamdia quelen (Hernandez et
al., 2009. The thicker of the muscularis layer is
related with the storage and expulsion of fecal
material from the intestine (Grau et al., 1992).
Proximal region of intestine in Clarias
batrachus showed higher amount of folds

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increasing the absorptive surface and

enhancing the absorptive activity. This is a
structural arrangement of the omnivorous
feeding habit by increasing efficiency,
whenmotility is increased. The complex folding
of intestinal mucosa with increased surface
area aids in mixing of food with hepatic,
pancreatic juices as well as with mucous
secreted by goblet cells. Similar results and
finding are reported in Mugil cephalus (ElBakary and El-Gammal 2010), in Seriola
dumerili (Grau et al., 1992) and in teleostean
(Khanna, S.S. 1971). The amount of mucosal
fold in the middle and distal region were found
to be less as compared to proximal region in
Clarias batrachus.
The intestine of Clarias batrachus contains
goblet cells in the mucosa of proximal, middle
and distal intestine. The population of goblet
cells increases toward the distal intestine
observed in the present study. This result of
increasing number of goblet cells is in
accordance with the reports in fish such as rice
field eel and flower fish Pseudophoxinus
antalyae (Cinar and Senol, 2006). The
increase amount of goblet cells may be
required for increased mucosal safety and
lubricant for faecal exclusion (Dai et al., 2007).
These goblet cells react positively with PAS
and AB, representing that they contain neutral
as well as acidic glycoproteins (Cinar and
Senol, 2006; Sucmez and Ulus, 2005, Bucke,
1971
and
Radaelli
2000).
Neutral
mucosubstances combine with alkaline
phosphate involve in emulsification of food into
chyme in vertebrates. Acidic mucins act to
protect the intestinal epithelium against the
enzyme glycosidase (Carrasson et al., 2006).
Thus, the present investigations of histological
studies of the intestine were similar to those
reported of other teleost, cat fishes and also
vertebrates. However there were some
variations observed in the presence of gastric
glands in pyloric region of stomach and goblet
cells were abundant towards the posterior
intestinal part of Clarias batrachus. The
histochemical finding showed positive for PAS
and AB in the intestinal region.
B. Ultrastructural study:
According to its feeding habits Clarias

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Histology, histochemical and ultrastructural studies on Intestine
of Indian catfish, Clarias batrachus (Linn 1758)

batrachus may consider as an omnivorous cat

fish. Variation in the gut region is found
generally in omnivorous fishes (Kapoor et al.,
1975). The ultrastructural study of intestine
show the epithelial lining of absorptive cell
from proximal intestine contains the usual
organelles and the regular microvilli which
increase the surface area of the intestine. The
cytoplasmic channels, lysosomal system,
microtubules, small vesicles (pinocytotosis)
are typical features of the lining cells of the
proximal, middle as well as distal intestine
region. In this study, the presence of microvilli,
tight junctional complexes, lysosome and
mitochondria in the enterocytes, are similar to
those observed in the freshwater fishes
(Kuperman and Kuzmaina, 1994), in Solea
solea (MacDonald, 1987), in Tilapia spp.
(Gargiulo, et al., 1998), in Solea senegalensis
(Arellano, 2002), in Orthrias angorae (Sucmez
and Ulus 2005) and in Oncorhynchus mykiss
(Banan Khojasteh et al., 2009).
The presence of the pinocytotic inclusion
provides evidence that an intracellular protein
digestion in the intestinal region may be
important in the fish species (Gargiulo et al.,
1998). Another feature of the mucosa of the
proximal, middle as well as the distal intestine
of C. batrachus is the presence of the lamellar
structure in the cytoplasm. This structure is
similar to those found in the cytoplasm
detected in the Senegal sole intestine
(Arellano, 2002). While, they were extreme in
the intestine of different larvae as well as adult
fishes (Noaillac-Depyre and Gas, 1979;
Calzada, et al., 1998). The lamellar structures
increase the surface area of membrane that
contact with the extracellular spaces maybe
facilitating
lipoprotein
transport.The
contribution of these intestinal membrane
infoldings in the transport of lipoprotein has
been recommended (Deplano et al., 1991).
While in Sparus aurata, they are associated
with the mitochondria in the basal cytoplasm
(Calzada et al., 1998). Moreover occurrences
is probably due to increase demand of energy
for process of osmoregulation, since the gut is
the main organ for absorbing the water for
maintain the water as well as ion balance in
case of marine fish species (Tytler and Ireland,

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1994)
The endocrine cells are restricted to the
proximal intestine (Gargiulo et al., 1998)
whereas in the present study the endocrine
cells are found in the proximal, middle as well
as in the distal intestinal region in C.
batrachus. We are first to report, the presence
of endocrine cells in the intestine of C.
batrachus. On the basis of the present
ultrastructural
result
three
types
of
enteroendocrine cells can be distinguish in the
proximal, middle as well as the distal intestine
of C. batrachus. Mainly these cells are of the
open type. This kind of cells may be
receptor-sensory, receiving specific chemical
information from top so as to stimulate or
inhibits the secretion at the base (Fujita and
Kobayashi, 1974). This hypothesis is
supported by the presence of microvilli, small
vesicles
(pinocytosis)
and
longitudinal
microtubules in the apical end of the endocrine
cell. These features are also described in
receptor cells of taste buds (Murray, 1973).
Some closed types of endocrine cells are also
found in the middle and proximal intestine.
Fujita and Kobayashi reported that this cell
type stimulate by the physiological stimuli
(Pressure, tension and temperature). Both
open and closed type of cells may be receiving
humoral stimuli from the bloodstream that
influence the release of granules.
In conclusion, the histological, histochemical
and ultrastructural feature of the proximal
middle and the distal intestine of Clarias
batrachus revealed an adaptation for the
feeding habits and this is important for the
protection and increasing of the absorptive
processes.

Acknowledgment: We are special thanks to

Dept. of Anatomy, All India Institute of Medical
Science, New Delhi, (INDIA) for providing
Electron Microscope Facility.

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