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Functional Zonation of Different Digestive Enzymes in

Etroplus suratensis and Oreochromis mossambicus
Hari Sankar. H.Sa&b, Jisha Joseb, Remya Varadarajanb, Smitha. V. Bhanub, Susan Joyb, Babu Philipb
a
Inter University Center for Development of Marine Biotechnology, Cochin University of Science and Technology, Fine Arts Avenue, Cochin-16,
Kerala, India
b
Department of Marine Biology, Microbiology and Biochemistry, School of Marine Sciences, Cochin University of Science and Technology, Fine
Arts Avenue, Cochin-16, Kerala, India

Abstract- The major digestive enzyme activities and digestive matching of an artificial diet to their nutritional needs [5]. The
indices were compared between Etroplus suratensis and understanding of the functional properties and optimal conditions
Oreochromis mossambicus. Pepsin-like acid proteases that acts for hydrolysis of nutrients by digestive enzymes in fish will
on low pH has been identified all along the digestive tract of both facilitate a more precise measurement of nutrient digestibility by
the fishes. Comparatively low alpha amylase activity is shown by a particular species. Fish can vary their feed from plankton in the
the E. suratensis and the enzyme is distributed almost equally summer to fish in the winter, occasionally even exploiting the
throughout the intestinal segments in both the species. Very low bacteria and algae of the water as a source of food [6]. Digestive
alkaline protease activity is found in the stomach of both the tissues are notoriously plastic in their responses to dietary change
fishes and in O. mossambicus, the enzyme activity diminishes [7]. Albeit of the food habit, the adaptations of the digestive
extensively towards the posterior portion of the intestine whereas system of different fish species exhibit closer correlation with
in E. suratensis the activity increases towards the posterior part. their diet than on their microenvironment and taxonomic
The present study showed that lipase is one of the prominent category [8]. Several researchers identified variations in gut
digestive enzymes in O. mossambicus with a remarkable specific morphology in response to fasting, increases in food intake and
activity throughout the digestive tract than that of E. suratensis. changes in diet [7, 9]. Thus, diet is a strong predictor of both
It has been noted that O. mossambicus has a higher values for intra- and inter-specific variation in the intestinal length,
digestive somatic index, hepato somatic index, intestinal indicating that fish adjust their phenotype to balance nutritional
coefficient and gut Vs standard length ratio than that of E. needs against energetic costs [10]. Therefore, it is necessary to
suratensis indicating its higher digestive and metabolic analyze some morphometric parameters of the digestive tract,
capabilities. The early maturity and fast growth of O. such as intestinal coefficient, digestive somatic index, and
mossambicus can be explained by their enhanced digestive hepatosomatic index of the selected teleosts. The aim of the
indices. The comparatively low activities of acid protease, present work is to study about the acid protease, amylase, lipase
amylase, lipase and total alkaline protease of E. suratensis and total alkaline protease digestive enzymes of Etroplus
revealed poor digestive capacity than that of O. mossambicus. suratensis and to compare with that of Oreochromis
mossambicus in order to increase our knowledge on the digestive
Index Terms- A Etroplus suratensis; Oreochromis mossambicus; physiology of these cichlid species and gain information
Digestion; Enzyme; Fish; Functional zonation concerning its nutrition.

I. INTRODUCTION II. MATERIALS AND METHODS

E troplus suratensis, (Pearl Spot, Karimeen), generally

considered as the ‘upper-middle class' fish has got a profile
uplift as the official state fish of Kerala state in India because of
Fish and preparation of crude enzyme extract
Experimental fishes of almost similar size (10-12cm) have
been collected from the Fisheries station, Kerala University of
its economical importance. The knowledge about digestive Fisheries and Ocean Studies, Puthuvyppu, India. The fishes were
physiology of E. suratensis is very limited. Tilapias are one of acclimated to laboratory condition for a week. A commercial diet
the most important euryhaline finfish cultured allover the world with known proximate composition has been given ad libitum.
and they represent approximately 6% of total farmed fish The fishes were starved for approximately 12 h prior to
production [1]. Oreochromis mossambicus is one of the hardiest sampling, subsequently killed by cold shock, and dissected
fishes in aquaculture farms. Once introduced into a habitat, they immediately. Among bony fishes, the pancreatic tissue is usually
generally establish themselves very quickly. The digestive diffused in or around the liver [11]. The exocrine pancreatic
enzymes play an important role in the development and growth tissue of O. niloticus has a diffused distribution in the hepatic
of fishes. The ability of the fish to utilize ingested nutrients parenchyma and is separated from the hepatocyte cords by means
depends on the activities of digestive enzymes present in various of thin septa of connective tissue [12]. Thus, in the present study
locations along the digestive tract. the whole liver consisting of diffused pancreatic tissue
The polyculture of different species of fish is an accepted (hepatopancreas) and the stomach was taken the as digestive
practice to promote fish production in India. The studies on organ. The stomach contents were squeezed out and rinsed with
digestive proteases of several species helped in the development cold distilled water to remove feed remnants. Since intestine of
of cost effective diets for their intensive farming [2, 3, 4] and the tilapia and pearlspot lacked visually distinct regions, they were

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 569
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divided into three segments of equal length and designated as III. RESULTS
anterior, middle and posterior intestine without squeezing or Pepsin like acid proteases activity has been observed all
rinsing. Ten percent (w/v) tissues homogenate was prepared in along the digestive tract of both E. suratensis and O.
cold Tris–HCl 50 mM buffer pH 7.2 using an electric mossambicus (Fig. 1). Considerably high acid protease activity
homogenizer (KEMI Model No: KHH 1), in ice-cold condition. has been established by O. mossambicus than that of E.
The homogenate was then centrifuged at 4°C at 10,000g for 10 suratensis and the activity decreases gradually towards the
min. The supernatant containing the enzymes was stored at -200C posterior part of the intestine. In comparison with stomach acid
until the analysis. protease activity in E. suratensis approximately two-fold increase
has been observed in O. mossambicus. Comparatively low alpha
Methods amylase activity is shown by the E. suratensis and the enzyme is
The tissue homogenates were purified by Trichloroacetic acid distributed almost equally through out the intestinal segments in
(TCA) precipitation and the precipitate of soluble protein was both the species. However, the middle intestine showed an
dissolved in 0.1M NaOH. The soluble protein content of enzyme increased alpha amylase activity (Fig. 2). Very low alkaline
extract was measured in comparison with BSA standards [13] by protease activity is found in the stomach of both the fishes. The
using Hitachi-2900 UV-Visible spectrophotometer. enzyme activity diminishes intensively at the posterior intestine
a) Acid Protease in O. mossambicus but in E. suratensis the activity extends until
Pepsin like acid protease was determined by Anson’s method the end of digestive tract (Fig.3) and the posterior part showed its
[14] and the specific enzyme activity is expressed in Anson Unit. maximum activity. The present study shows that lipase is one of
b) Total alkaline protease the prominent digestive enzymes in O. mossambicus with a
The total alkaline protease was estimated by using Casein as remarkable specific activity throughout the digestive tract (Fig.
substrate [15] with a slight modification in tris buffer pH as 8. 4). Very low activity of stomach lipase is shown by E. suratensis
Briefly, 50µl of homogenate was mixed with 0.1M Tris buffer (Fig. 4) compared to O. mossambicus. The results have been
pH-8 having 20mM CaCl2 and preincubated for 5 minutes at expressed as bar diagrams that represents mean ± standard
370C and mixed thoroughly with 1% substrate solution. The deviation. Each set of bars values with different lower case
reaction was stopped by adding 12% ice cold TCA and tyrosine letters vary significantly (p<0.05) in each tissue on the two
in the supernatant was measured at 280nm after centrifugation at different species (One-way ANOVA).
8000 rpm for 15 min. The homogenate was added to the blank The average length and weight of fishes showed very good
tubes at the end of the incubation after adding the TCA. L- correlation (0.99) in the case of E. suratensis but in the case of O.
tyrosine was used as a standard, and one unit of enzyme activity mossambicus it is not that good (0.73) (Table 1). O.
(U) is defined as the amount of enzyme needed to catalyze the mossambicus has a higher index for DSI, HSI, Intestinal
formation of µM of tyrosine/min/ ml of homogenate at 37 0 C. coefficient and gut Vs standard length ratio than that of E.
The specific activity is expressed as U/mg protein. suratensis (Table 1).
c) Lipase
Lipase was estimated spectrophotometrically by hydrolysis of
p-nitrophenyl palmitate using a modified method based on that of IV. DISCUSSION
Winkler and Stuckmann [16]. 50µl of 20mM p-nitrophenyl
palmitate was mixed with 20mM Tris buffer (pH-8) containing Etroplus suratensis is the prime among the cichlids
20mM CaCl2, 5mM sodium cholate and 0.01% gum Arabica. indigenous to peninsular India and Srilanka. It is one of the most
50µl of tissue homogenate was added to this reaction mixture popular and promising species for aquaculture in India because
and incubated for 10 minutes. The p-nitro phenol liberated was of its high market demand and large size. With the flourishing of
read at 410nm against a reagent blank. backwater tourism in Kerala, the demand for Pearl Spots has
d) Amylase been on the increase. However, this species is facing serious
The reducing sugars liberated by the action of alpha- amylase diminution in its natural habitats owing to unrestrained
on starch was estimated by Somogyi–Nelson method using 3,5- exploitation and challenge by invasive species like tilapia. The
dinitrosalycylic acid (DNSA) [17]. One unit of activity (U) is general trend to exploit the most valuable species increased the
defined as the amount of enzyme able to produce 1 mg of threat in its maximum. It is highly nutritive, besides a good
maltose/min/ml of homogenate at 370 C. The specific activity is amount of meat, pearlspot include protein (16.74 %), lipids (1.15
expressed as U/mg protein. %), carbohydrates (2.43 %), moisture (78.55%), Zn (83.04μg/g),
Zootechnical indices Mn (39.44μg/g), Mg (1.15mg/g), Co (8.56μg/g), Cr (33.96μg/g),
The indices are calculated as following Cu (24.92μg/g), Fe (566.40μg/g), Cd (1.12μg/g) and Ni
Digestive somatic index (DSI) = (Digestive tract (2.64μg/g) [18]. Experimental information on the metabolic and
weight/Body weight) X 100 digestive enzyme profile of Pearl Spot is inadequate for the
Hepatosomatic index (HSI) = (Hepatopancreas weight/Body formulation of efficient compound feeds.
weight) X 100 Distribution and activity of intestinal digestive enzymes
Intestinal coefficient = Digestive tract length/Total fish along the intestinal tract varies with feeding habit and intestinal
length morphology [19-22]. The reports on digestive physiology of E.
Intestinal to Standard Length ratio = Total gut length/ Standard suratensis were very limited. In Chinook salmon, weight gain
length was found to be positively correlated with the ability of the
digestive enzymes to hydrolyze diets [23]. In this study, it is

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found that proteases are very prominent in both stomach and could be the reason for a slightly increased alpha amylase
intestinal segments and they are widely distributed though out activity in that segment.
the alimentary canal. Various studies on other fish digestive A comparative study of the activity of digestive proteolytic
secretions have shown the occurrence of acid proteases that have enzymes and amylase could reveal the capacity of different
high activity in the acidic region in the stomach and alkaline species to use protein and carbohydrates [36]. It was adumbrated
proteases acting actively in alkaline pH region in the intestine that changes in digestive enzyme activity could be affected by
[24, 25]. Proteolytic activities at low pH have also been reported feeding behavior and biochemical composition of food [40]. The
in species with prominent stomach region and a high pepsin adaptations of the digestive system of different species exhibit
secretion such as eel, tilapia, salmon, sea bass and trout [25, 26, closer correlation with their diet rather than on their taxonomic
27, 28, 29]. Pepsin has been identified as the first proteolytic category [8]. Antithetically, there are studies on the phylogeny
enzyme acting in fish digestive tract as a major acidic protease which influences the pattern of amylase activity more than that
[20, 30]. Pepsin, a member of the aspartic endopeptidase family, by diet in prickleback fishes [41]. Earlier reports on E. suratensis
has been identified in several species [23]. In the present indicated that the young ones of this species are herbivores [42,
examination, the acid protease such as pepsin is found to be 43]. It has been observed that though it feeds on micro and macro
present in all segments of the alimentary tract of both E. vegetation, invertebrates such as insect larvae, bivalves, mysids
suratensis and O. mossambicus. It has been revealed that the and decayed organic matter are mainly consisted in its food [42,
peptic digestion in Tilapia nilotica is absent [31]. However, in 43, 44, 45]. Carbohydrases and proteolytic activities were higher
the natural physiology of digestion, the acid protease will not be in the detritivore compared to the omnivorous and carnivorous
active in the intestinal segments due to the neutralization of fishes [46]. In contrary, the present study obtained comparatively
acidic condition by the action of bile. For proper utilization of higher α-amylase and both acid and alkaline protease activity in
proteins, tilapia requires a highly acidic medium to enable O. mossambicus than that of E. suratensis.
biochemical digestion of protein due to thin stomach walls when The carnivorous fishes have higher lipase activity compared
compared to fishes with muscular stomach like African catfish, to herbivorous and omnivorous fishes [30, 47] and it is attributed
which relies more on mechanical breakdown of nutrients and by the higher consumption of fat rich food by carnivorous fishes
possesses lower pepsin secretion [32]. [35]. The lipase is not necessarily produced by the pancreas but
After gastric digestion, the protein digestion is completed by is a property of hepatic tissue [6]. Lipolytic activity in fish is
the basic proteases of intestinal and pancreatic origin. A high generally higher in the proximal part of the intestine and the
activity of basic protease enzyme activity is observed in both pyloric caeca, if present. It can extend up to the lower parts of the
species. Enzymes such as trypsin, chymotrypsin, collagenase, intestine with the activity decreasing progressively. Low lipolytic
elastase, and carboxypeptidase have been characterized in activity has also been found in the stomach of several fishes but
different types of fish [33, 34]. Precursor zymogens of the the physiological significance of gastric lipolytic activity in fish
alkaline proteases are secreted by pancreas [35]. High protease is unclear [48]. The pancreas or hepatopancreas is generally
activity by intestinal extract at different alkaline pH range has considered as the major source of digestive lipase enzymes in
been shown by different researchers in various species like carp fish as it is in mammals [37, 49]. The lipolytic activity found in
[26], rainbow trout and Atlantic salmon [28], halibut and turbot stomach is different from that of pancreatic origin, suggesting
[4], striped and European sea bass [33], sea bream and dentex that stomach may be a source of lipases, and the intestinal flora
[3], goldfish [36] and discus fish [24]. It provides evidence for also contributes lipolytic activities in the digestive tract of fish
the presence of minimum two major groups of alkaline proteases [48]. In the present study O. mossambicus possesses a high
with different optimum pH. The E. suratensis has a short and less profile of lipase activity. It should be helpful in the effective
coiled intestine and the alkaline protease activity that gradually digestion of lipids available in the various feeds of plant or
increases along the digestive tract reaches its maximum at the animal origin. Comparatively a low lipase profile has been
posterior intestine. O. mossambicus has a highly coiled, thin and exhibited by E. suratensis. The anterior and middle part of both
elongated intestine with a maximal activity at the middle the fishes showed the most intense activity of lipase similar to
segment. the results similar to previous studies [30].
In comparison to proteases, knowledge about carbohydrases In teleosts, the gastrointestinal tract morphology generally
and lipases are still lacking in many species, despite the reported shows specific variations with respect to diet, feeding habit
importance of these enzymes [36]. Various workers have phylogeny, body shape, and features that reflect functional
demonstrated that amylase activity is greater in omnivorous and differentiation [49, 50]. In the present study, O. mossambicus has
herbivorous fish than in carnivorous fish [36, 37, 38]. Low or a higher index for DSI, HSI, intestinal coefficient and gut Vs
moderate amylase activities have been reported in other standard length ratio than that of E. suratensis indicating higher
carnivorous species [39]. Comparatively a lower activity of digestive and metabolic capability. The fast growth and attaining
amylase is detected in the stomach in this study and it has of maturity of O. mossambicus can be explained from the
previously been suggested that the presence of amylase in differences in digestive indices. In addition, there is a correlation
stomach could be due to some exogenous contamination from between the structure of the digestive apparatus and the feeding
intestinal activity [25, 32]. In the natural environment, habit of fishes [51, 52]. It is commonly emphasized that
carbohydrates are indeed more predominant than protein. Thus, herbivore and detritivore fish species tend to have longer, thinner
there is possibility of carbohydrate digestion beginning from the and narrower intestines than carnivores [37, 51] and the intestine
stomach [20, 39]. The neutralization of chime by bile may not be of omnivorous species have an intermediate length [52].
complete before it reaches the middle intestinal segment and this

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V. CONCLUSION [12] C. A. Vicentini, I. B. Franceschini-Vicentini, M. T. S. Bombonato, B.

Bertolucci, S. G. Lima, A. S. Santos, Morphological study of the liver in the
The comparative study on digestive enzymes such as acid teleost Orechromis niloticus. Int J Morphol, Vol. 23, 2005, 211-216.
and alkaline protease, amylase and lipase activity in E. suratensis [13] O. H. Lowry, N. J. Rosebrough, A. L. Farr, R. J. Randall, Protein
and O. mossambicus, shows that O. mossambicus would digest measurement with folin-phenol reagent, J Biol Chem, Vol. 193, 1951, 265–
275.
the dietary proteins and fats better than E. suratensis and its
[14] M. L. Anson, The estimation of pepsin, trypsin, papain and cathepsin with
amylase pool would enable this fish to digest carbohydrates at haemoglobin. J Gen Physiol, Vol. 20, 1938, 79-89.
herbivorous levels. When compared, O. mossambicus exhibit the [15] L. M. Lundstedt, J. F. B. Melo, G. Moraes, Digestive enzymes and
omnivorous gut characteristics like long and highly coiled metabolic profile of Pseudoplatystoma corruscans (Teleostei: Siluriformes)
intestine, but E. suratensis has a short intestine. The knowledge in response to diet composition. Comp Biochem Physiol, Vol. 137 B, 2004,
about the digestive enzymes and digestive capabilities would be 331–339
advantageous for diet manufacturing, as carbohydrates could be [16] . K. Winkler, M. Stuckmann, Glycogen, hyaluronate, and some other
polysaccharides greatly enhance the formation of exo-lipase by Serratia
added at a greater proportion than protein and thereby save on marcescens. J Bacteriol, Vol. 138, 1979, 663–670
feed manufacturing costs. In turn, the higher proportion of [17] P. Bernfeld, Amylases, α and β: colorimetric assay method, in Methods in
protease enzymes and amylase with respect to lipase in pearlspot Enzymology edited by S P Colowick and N O Kaplan (Academic Press,
may help to reduce the fat content in feed, thereby increase in New York), Vol.1, 1955, 149-158.
shelf life of artificial feed for this species. This knowledge will [18] S. Shanker, G. Marichamy, A. Saradha, A. R. Nazar, M. A. Badhul Haq,
be helpful to understand the competitive feeding strategies of Proximate composition and bioaccumulation of metals in some fin fishes
and shellfishes of Vellar Estuary (South east coast of India). Euro J Exp
native and alien species. Bio, Vol. 1, 2011, 47-55.
[19] V. V. Kuz’mina, Relative enzyme activity in the intestinal lumen and
mucosa. J Ichthyol, Vol. 24, 1984, 140-144.
ACKNOWLEDGEMENTS [20] U. Sabapathy, L. H. Teo, A quantitative study of some digestive enzymes in
the rabbitfish, Siganus canaliculatus and the sea bass Lates calcarifer, J Fish
The authors are thankful to the Head, Department of Marine Biol, Vol. 42, 1993, 595–602.
Biology, Microbiology and Biochemistry, CUSAT for providing [21] S. Kolkovski, Digestive enzymes in fish larvae and juveniles-Implications
facilities for conducting this work. We express our gratefulness and applications of formulated diets. Aquaculture, Vol. 200, 2001, 181-201.
to Prof. (Dr) M.M. Jose, Fisheries Station, Kerala University of [22] A. Gawlicka, S. S. O. Hung, M. A. Herold, F. T. Barrows, J. de la Noue ,
Fisheries and Ocean Studies, Puthuvyppu, Kerala, India for the Effects of dietary lipids on growth, body fatty acid composition and tissue
enzyme histochemistry of white sturgeon, Acipense transmontanus larvae. J
supply of E. suratensis and O. mossambicus. Appl Ichthyol, Vol. 18, 2002, 673-681.
[23] N. F. Haard, L. E. Dimes, R. E. Arndt, F. M. Dong, Estimation of protein
digestibility: IV. Digestive proteinases from the pyloric caeca of coho
REFERENCES salmon Oncorhynchus kisutch fed diets containing soybean meal. Comp
Biochem Physiol, Vol. 115B, 1996, 533–540.
[1] FAO (Food and Agriculture Organization of the United Nations), Fishstat
[24] A. S. C. Chong, R. Hashim, L. Chow-Yang, A. B. Ali, Partial
Plus., Aquaculture production, 2004, 1950-2002.
characterization and activities of proteases from the digestive tract of
[2] J. Clark, N. L. MacDonald, J. R. Stark, Metabolism in marine flatfish. II. discus fish (Symphysodon aequifasciata). Aquaculture, Vol. 203, 2002,
Protein digestion in Dover sole (Solea solea L.). Comp Biochem Physiol, 321-333.
Vol.81B, 1985, 217-222.
[25] D. M. Xiong, C. X. Xie, H. J. Zhang, H. P. Liu, Digestive enzymes along
[3] F. J. Alarcon, M. Diaz, F. J. Moyano, E. Abellan, Characterization and digestive tract of a carnivorous fish Glyptosternum maculatum (Sisoridae,
functional properties of digestive proteases in two sparids; gilthead Siluriformes). J Anim Physiol Anim Nutr, Vol. 95, 2011, 56–64.
seabream (Sparus aurata) and common dentex (Dentex Dentex). Fish
[26] E. Jonas, M. Ragyanssszki, J. Olah, L. Boross, Proteolytic digestive
Physiol Biochem, Vol.19, 1998, 257–267.
enzymes of carnivorous Silurus glanis L., herbivorous
[4] H. J. Glass, N. L. MacDonald, R. M. Moran, J. R. Stark, Digestion of Hypophthalmiichtthys molitrix Val. and omnivorous Cyprinus carpio L.
protein in different marine species. Comp Biochem Physiol, Vol. 94B, fishes. Aquaculture, Vol. 30, 1983, 145–154.
1989, 607–611.
[27] S. S. Twining, P. A. Alexander, K. Huibregste, D. M. Glick, A pepsinogen
[5] M. Furne, M. C. Hidalgo, A. Lo´pez, M. Garcıa-Gallego, A. E. Morales, A. from rainbow trout. Comp Biochem Physiol, Vol. 75B, 1983, 109 – 112.
Domezain, J. Domezaine, A. Sanz, Digestive enzyme activities in Adriatic
[28] K. R. Torrissen, Characterization of proteases in the digestive tract of
sturgeon Acipenser naccarii and rainbow trout Oncorhynchus mykiss. A
Atlantic salmon Salmo salar in comparison with rainbow trout Salmo
comparative study. Aquaculture, Vol. 250, 2005, 391– 398.
gairdneri. Comp Biochem Physiol, Vol. 77B, 1984, 669–674.
[6] E. J. W. Barrington, The alimentary canal and digestion, in The Physiology
[29] A. Yamada, K. Takano, I. Kamoi, Purification and properties of protease
of Fishes edited by M E Brown (Academic Press, New York), 1957, 109-
from tilapia stomach. Nippon Suisam Gakkaishi, Vol. 59, 1993, 1903–
161.
1908.
[7] J. M. Starck, Structural flexibility of the gastro-intestinal tract of
[30] B. Tengjaroenkul, B. J. Smith, T. Caceci, S. A. Smith, Distribution of
vertebrates: implications for evolutionary morphology. Zoologischer
intestinal enzyme activities along the intestinal tract of cultured Nile tilapia,
Anzeiger, Vol. 238, 1999, 87 - 101.
Oreochromis niloticus L. Aquaculture, Vol. 182, 2000, 317–327.
[8] I. Fernandez, F. J. Moyano, M. Diaz, T. Martinez, Characterization of α-
[31] D. J. W. Moriarty, C. M. Moriarty, The assimilation of carbon from
amylase activity in five species of Mediterranean sparid fishes (Sparidae,
phytoplankton by two herbivorous fishes, Tilupiu nilotica and
Teleostei). J Exp Mar Biol Ecol, Vol. 262, 2001, 1-12.
Haplochromis nigripinnus, J Zool, Vol.171, 1973, 41-55.
[9] J. Olsson, M. Quevedo, C. Colson, R. Svanback, Gut length plasticity in
[32] W. Uys, T. Hecht, Assays on the digestive enzymes of sharptooth catfish,
perch: into the bowels of resource polymorphisms. Biol J Linn Soc, Vol. 90,
Clarias gariepinus (Pisces: Clariidae). Aquaculture, Vol. 63, 1987, 303–313.
2007, 517–523.
[33] A. Eshel, P. Lindner, P. Smirnoff, S. Newton, S. Harpaz, Comparative study
[10] C. E. Wagner, P. B. McIntyre, K. S. Buels, D. M. Gilbert, E. Michel, Diet
of proteolytic enzymes in the digestive tracts of the European sea bass and
predicts intestine length in Lake Tanganyika’s cichlid fishes. Funct Ecol,
hybrid striped bass reared in freshwater. Comp Biochem Physiol, Vol.
Vol. 23 2009, 1122–1131.
106A, 1993, 627–634.
[11] C. E. Bond, Biology of Fishes (Saunders College Publishing, Philadelphia),
1979, 38.

www.ijsrp.org
International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 572
ISSN 2250-3153

[34] S. T. Chiu, B. S. Pan, Digestive protease activities of juvenile and adult ee1 Third Author – Remya Varadarajan, Department of Marine
(Anguilla japonica). Aquaculture, Vol. 205, 2002, 141– 156.
Biology, Microbiology and Biochemistry, School of Marine
[35] I. Chakrabarti, M. A. Gani, K. K. Chaki, R. Sur, K. K. Misra, Digestive
enzymes in 11 freshwater teleost fish species in relation to food habit and
Sciences, Cochin University of Science and Technology, Fine
niche segregation. Comp Biochem Physiol, Vol. 112A, 1995, 167–177. Arts Avenue, Cochin-16, Kerala, India
[36] M. C. Hidalgo, E. Urea, Sanz, Comparative study of digestive enzymes in Fourth Author – Smitha. V. Bhanu, Department of Marine
fish with different nutritional habits. Proteolytic and amylase activities. Biology, Microbiology and Biochemistry, School of Marine
Aquaculture, Vol. 170, 1999, 267–283. Sciences, Cochin University of Science and Technology, Fine
[37] R. Fange, D. Grove, Digestion, in Fish physiology, edited by W. S. Hoar, Arts Avenue, Cochin-16, Kerala, India
D. J. Randall & J. R. Brett, (Academic Press, New York), Vol. VIII, 1979,
pp 61–260. Fifth Author – Susan Joy, Department of Marine Biology,
[38] A. M. Ugolev, V. V. Egorova, V. V. Kuźmina, A. A. Grudskov, Microbiology and Biochemistry, School of Marine Sciences,
Comparative-molecular characterization of membrane digestion in fish and Cochin University of Science and Technology, Fine Arts
mammals. Comp Biochem Physiol, Vol. 76B, 1983, 627–635. Avenue, Cochin-16, Kerala, India
[39] R. Munilla-Moran, F. Saborido-Rey, Digestive enzymes in marine species: Sixth Author – Babu Philip, Department of Marine Biology,
II. Amylase activities in gut from seabream Sparus aurata, turbot Microbiology and Biochemistry, School of Marine Sciences,
Scopthalmus maximus and redfish Sebastes mentella. Comp Biochem
Physiol, Vol. 113B, 1996, 827–834. Cochin University of Science and Technology, Fine Arts
[40] V.V. Kuz’mina, I. L. Golovanova, G. I. Izvekova, Influence of temperature Avenue, Cochin-16, Kerala, India
and season on some characteristics of intestinal mucosa carbohydrases in
six freshwater fishes. Comp Biochem Physiol, Vol. 113B, 1996, 255-260.
[41] A. S. Chan, H. M. Horn, K. A. Dickson, A. Gawlicka, Digestive enzyme
activities in carnivores and herbivores: comparisons among four closely
related prickleback fishes (Teleostei: Stichaeidae) from a California rocky
intertidal habitat. J Fish Biol, Vol. 65, 2004, 848-858.
[42] K. H. Alikunhi, Fish culture in India. Fann Bull. Indian Coun Agne Res,
Vol. 20, 1957, 144.
[43] S. L. Hora, T. V. R. Pillay, Hand-book on the fish culture in the Indo-
Pacific region. FAD Fish Biol Tech Pap, Vol. 14, 1962, 124.
[44] R. D. Prasadam, Observation on the biology of the pearl-spot Etroplus
suratensis (Bloch) from the Pulicat lake, Madras, J Inland Fish Soc India,
Vol. 3, 1971, 72-78.
[45] Keshava, P. Santha Joseph, M. Mohan Joseph, Feeding habits of the Pearl-
Spot Etroplus suratensis (Bloch) in the Nethravati - Gurpur estuary. The
First Indian Fisheries Forum. Proceedings, 1988, 203-206.
[46] K. Lopez-Vásquez, C. A. Castro-Pérez, A. L. Val, Digestive enzymes of
eight Amazonian teleosts with different feeding habits, J Fish Biol, Vol. 7,
2009, 1620-1628.
[47] K. Opuszynski, J. V. Shireman, Digestive mechanisms. In Herbivorous
Fishes: Culture and Use for Weed Management edited by K. Opuszynski, J.
V. Shireman (CRC Press, Boca Raton, FL), 1995, 21– 31.
[48] R. E. Olsen, E. Ringo, Lipid digestibility in fish: a review. Recent Res Dev
Lipids Res, Vol. 1, 1997, 199–265.
[49] B. C. Kapoor, H. Smith, I. A. Verighina, The alimentary canal and digestion
in teleosts. Adv Mar Biol, Vol. 13, 1975, 109–239.
[50] M. A. Abaurrea, M. I. Nuikiez, M. V. Ostos, Ultra structural study of the
distal part of the intestine of Oncorhynchus mykiss. Absorption of dietary
protein. Micron, Vol. 24, Issue 5, 1993, 445-450.
[51] R. Fugi, A. A. Agostinho, N. S. Hahn, Trophic morphology of five benthic-
feeding fish species of a tropical floodplain. Braz J Biol, Vol. 61, 2001, 27-
33.
[52] B. M. S. Ward-Campbell, F. W. H. Beamish, C. Kongchaiya,
Morphological characteristics in relation to diet in five coexisting Thai fish
species. J Fish Biol, Vol. 67, 2005, 1266-1279.

AUTHORS
First Author – Hari Sankar. H.S, Inter University Center for
Development of Marine Biotechnology, Cochin University of
Science and Technology, Fine Arts Avenue, Cochin-16, Kerala,
India
Second Author – Jisha Jose, Department of Marine Biology,
Microbiology and Biochemistry, School of Marine Sciences,
Cochin University of Science and Technology, Fine Arts
Avenue, Cochin-16, Kerala, India

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 573
ISSN 2250-3153

Table 1: Length-Weight relationships and digestive indices of Etroplus suratensis and Oreochromis mossambicus

TL SL TW COR TGL TGW HW SW DSI HIS IC Gut Vs SL Ratio

E. suratensis 11.950±1.31 9.333±1.0 42.167±14.89 0.99 41.167±5.74 0.37±0.06 0.613±0.42 0.165±0.03 0.956±0.307a 1.332±0.475a 3.445±0.303a 4.411±0.391a

O. mossambicus 11.283±0.64 8.833±0.41 21.19±2.58 0.73 65.50±21.64 0.977±0.24 0.305±0.14 0.133±0.05 4.584±0.874b 1.475±0.701a 5.755±1.61b 7.353±2.11b

TL- Total Length, SL- Standard Length, TW- Total Weight, COR-Correlation between Total length and Total weight, TGL- Total Gut Length, TGW- Total Gut Weight, HW-
Hepatopancreas Weight, SW- Stomach Weight, IC- Intestinal coefficient. Length was measured in centimeter and weight in grams and they were reported as mean ± Standard
deviation (SD). Digestive somatic index (DSI), Hepato-somatic index (HIS), Intestinal coefficient and Gut Vs Standard Length Ratio were reported as mean ± SD. On each
columns values with different lower case letters vary significantly (P<0.05) in each tissue on the two different species (One-way ANOVA)

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 574
ISSN 2250-3153

Distribution of Acid Proteases

0.016 c Stomach
Hepatopancreas

Specific Activity in Anson U/mg

0.014
Anterior
0.012 M iddle
b
0.01 Posterior
b ab

Protein
0.008
a
0.006 a a
a a
0.004 b

0.002

0
Etroplus suratensis Oreochromis mossambicus
Species

Figure 1: Distribution of acid protease along the digestive segments of E. suratensis and
O. mossambicus

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 575
ISSN 2250-3153

Distribution of alpha-Amylase

1.2 c
c Stomach
d
1 Hepatopancreas
Anterior

Specific activity
0.8 d b M iddle
d
Posterior
0.6 c

0.4 b
a
0.2
a
0
Etroplus suratensis Oreochromis mossambicus
Species

Figure 2: Distribution of alpha-amylase along the digestive segments of E. suratensis and

O. mossambicus

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 576
ISSN 2250-3153

0.016 Distribution of Total Alkaline Protease

0.014 c
Stomach
0.012
Hepatopancreas

Specific Activity
0.01 d c
Anterior

0.008 M iddle
bc
0.006 bc ab
Posterior
b
0.004
ab
0.002 a a
0
Etroplus suratensis Oreochromis mossambicus
Species

Figure 3: Distribution of total alkaline protease along the digestive segments of

E. suratensis and O. mossambicus

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International Journal of Scientific and Research Publications, Volume 4, Issue 5, May 2014 577
ISSN 2250-3153

Distribution of Lipase Stomach

9 b
Hepatopancreas
8
Anterior
7 M iddle
a
a Posterior

Specific Activity
6

5 a
a
4

2 c
b b b
1 a
0
Etroplus suratensis Oreochromis mossambicus
Species

Figure 4: Distribution of lipase along the digestive segments of E. suratensis and

O. mossambicus

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