Advance Journal of Food Science and Technology 3(3): 160-164, 2011
ISSN: 2042-4876
© Maxwell Scientific Organization, 2011
Received: February 04, 2011
Accepted: March 08, 2011
Published: June 06, 2011
Studies on the Seasonal Variation in Lipid Composition of Muscles,
Hepatopancreas and Ovary of Freshwater Prawn, Macrobrachium dayanum
(Henderson) During Reproductive Cycle
Anshu Samyal, Yahya Bakhtiyar, Archana Verma and Seema Langer
P.G. Department of Zoology, University of Jammu, Jammu-180006, Jammu and Kashmir, India
Abstract: The Macrobrachium dayanum, a locally available freshwater prawn of Jammu was sampled regularly
for a period of one year to analyze the seasonal variation in the lipid contents of muscles, hepatopancreas and
ovary. Remarkable variation in the lipid content in different tissues of M. dayanum has been analyzed during
the study period. Changes in the lipid content in all the three tissues were observed to be statistically significant
(p<0.05). Lipids in muscles corresponds to 0.23±0.11 and 3.83±0.11%, minimum and maximum value in the
month of September and December respectively. In hepatopancreas minimum of 5.05±0.21% in August and
maximum of 9.31±0.41% in February have been observed. Mean value of lipid content in the ovaries range
from 6.14±0.24% (minima) to 9.76±0.80% (maxima) in July and April respectively. Moreover, maximum
numbers of stage IV oocytes are observed in the ovary in the months of March-April and August to October
accompanied with increase in the lipid content during these respective months.
Key words: Hepatopancreas, lipids, Macrobrachium dayanum, muscles, ovaries and Jammu
INTRODUCTION
MATERIALS AND METHODS
Prawns are very popular and sought for food all
around the world. Their palatability is closely linked with
their biochemical composition which changes
considerably with season and maturation process. Lipids
play significant role during the gonadal growth,
maturation and development of decapod crustaceans.
They are very important food reserves in the oocytes
(Gallager et al., 1986; Le Pennec et al., 1998).
Knowledge of the biochemistry and metabolism of the
processes that occur during maturation are essential for a
complete understanding of crustacean reproduction. Lipid
deposition during maturation is crucial to reproduction
and early larval development because lipids are known to
play several essential roles in the metabolism of
crustaceans. Apart from being a major source of
metabolic energy and the main form of energy storage,
lipids also supply essential fatty acids needed for the
maintenance and integrity of cellular membranes, and
serve as precursors of steroid and moulting hormones
(Middleditch et al., 1980; Harrison, 1990).
Keeping in view the essential role of the lipids an
attempt has been made to study the variation in the
quantity of lipids in muscles, hepatopancreas and ovary of
Macrobrachium dayanum. The data may lead to the better
understanding of the relative importance of the lipids
during reproductive cycle and during different seasons
of a year.
Collection: Prawns were collected from their natural
habitat from Gho-Manhasan stream located at a distance
of about 20 kms. North-West of Jammu city (32º67/ lat N;
74º79/ long E) during morning hours (800-1100 h). Live
specimens were then carried to the Gho-Manhasan fish
farm where they were oxygen packed in polythene bags
and brought to wet lab, Department of Zoology,
University of Jammu where they were kept in plastic
troughs. Male and female individuals in the size range of
4 to 5 cm above were segregated and kept in separate
troughs to analyze lipid content and different stages of
ovarian maturation.
Proximate body composition and microtomy: Monthly
collected samples of Macrobrachium dayanum were
subjected to biochemical analysis. This analysis was
performed over a period of one year, beginning in May
2006 and concluding in April 2007. Permanent slides
were prepared to study under different stages of ovarian
maturation by following standard methods of microtomy.
The slides so prepared were then studied under Nikon YS
100 microscope and photographed with the help of SDC313 Camera. Lipids in different tissues were estimated by
adopting the standard method of Folch et al. (1957).
Statistical analysis: The data was statistically analyzed
on personal computer to calculate correlation by
Corresponding Author: Yahya Bakhtiyar, P.G. Department of Zoology, University of Jammu, Jammu-180006, Jammu and
Kashmir, India.
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Adv. J. Food Sci. Technol., 3(3): 160-164, 2011
Table 1: Seasonal variation in the Lipid composition of Muscles,
Hepatopancreas, and Ovary of Macrobrachium dayanum
Months
Muscles
Hepatopancreas
Ovary
May
1.32±0.25ef
6.50±0.25c
9.05±0.51ab
June
2.76±0.11b
6.54±0.21c
7.28±0.32d
July
2.65±1.03bc
6.29±0.41c
6.14±0.24e
Aug
0.685±0.22fg
5.05±0.21e
8.91±0.45ab
Sept
0.23±0.11g
5.80±0.23d
9.24±0.25a
Oct
1.78±0.25de
7.35±0.32b
8.30±0.29bc
b
a
Nov
3.06 ± 0.46
9.11±0.42
7.85±0.25cd
Dec
3.83±0.11a
9.17±0.11a
7.46±0.56cd
Jan
2.06±0.11cd
9.23±0.17a
7.61±0.63cd
Feb
1.45±0.17de
9.31±0.42a
8.27±0.22bc
March
1.29±0.14ef
8.85±0.27a
8.96±0.58ab
April
1.78±0.25de
7.36±0.14b
9.76±0.80a
Data presented in the table is the mean of three readings i.e. Mean ± S.D
The values having the same superscript in a column do not differ
significantly
Pearson’s correlation method, ANOVA with the help of
MS Excel 2003 and SPSS (12.0 version Inc. Chicago,
USA) and mean compared by using Duncan’s multiple
range test taking p<0.005 as significance level
(Duncan, 1955).
RESULTS
Biochemical analysis of the muscles hepatopancreas and
ovaries of M. dayanum exhibited seasonal variation in
lipids. Changes in lipid composition concuss with changes
in season and reproductive activity (Table 1).
Muscles: During present studies lipid content of muscles
(Table 1) was observed to have a minimum value of
0.23±0.11% in September and maximum value of
3.83±0.11% in the month of December. An inquisitive
study also reveals that the muscle lipid content remains
characteristically low during post monsoon (August to
September) and maximum during winters (November to
January).
Hepatopancreas: Hepatopancreas (Midgut gland) is a
physiologically storage organ of organic reserve like
glycogen, lipids and protein. In M.dayanum, lipid content
in hepatopancreas varies from minimum of 5.05±0.21%
in August and maximum of 9.31±0.42% in February. It is
also evident from the recorded values (Table 1) that lipid
content undertakes a steady unabated fall from March to
September.
Ovary: Perusal of Table 1 reveals that mean value of
lipid content in the ovaries range from 6.14±0.24%
(minima) to 9.76±0.80% (maxima) in July and April
months respectively .The level of variations recorded in
Table 1 in ovaries of M. dayanum as depicted is very
small between 6.14±0.24 to 9.76±0.80%.
Four different stages of oocytes in the ovarian
development of the fresh water prawn, M. dayanum have
been recognized.
Fig. 1: Showing the stage I (Oogonia) of reproductive cycle of
Macrobrachium dayanum
Stage I: Oogonia (Fig. 1): In this stage, the oocyte is
either oval or spherical measuring about 20-52.0 : with
centrally located round nucleus. Nucleolus is also
observed either at the centre of nucleus or adhered to the
nuclear membrane. Cytoplasm is homogeneous without
any vacuoles or yolk globules. In few oogonia of this
stage binucleolate condition was also observed. During
the developmental process it is the first differentiable
oogonia and smallest in size. Oogonia are present either
attached to germinal epithelium or lie freely in the
ovocoel in the months of December to March and May to
July.
Fig. 2: Showing the stage II (Previtellogenic oocytes) of
reproductive cycle of Macrobrachium dayanum
seems to be homogenous but in certain cells it is subjected
to characteristic change with peripheral vacuolization i.e.,
vacuoles appear in the cytoplasm and are arranged
Stage II: Previtellogenic oocytes (Fig. 2): The oocytes
of this stage measure from 45.0-195.0 :. Ooplasm
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Adv. J. Food Sci. Technol., 3(3): 160-164, 2011
Stage IV: Mature oocytes (Fig. 4): Oocytes of this stage
attain maximum size of about 650-950 :. The yolk
globules occupy the entire ooplasm. Nucleus is
acentrically located. Yolk globules present at the
periphery are spherical in shape and those located
centrally are oval in shape and small sized. Each oocyte
is surrounded by a clear demarcated layer of follicular
epithelium. Between the follicular epithelium and the
inner ooplasm develops a thin egg membrane. Maximum
number of oocytes of this stage is present in the months of
March-April and August to October.
DISCUSSION
M. dayanum is a biannual breeder: first phase of
annual breeding occurs in the months of April-June and
second phase of annual breeding lasts from August to
September. A similar trend of biannual breeding in
crustaceans is already on record (Kailoo, 1984;
Malik, 2006). A large number of workers have estimated
the lipid content of muscles in decapod Crustaceans
(Chanmugam et al., 1983; Gopakumar, 1993;
Cavalli et al., 2001; Rosa and Nunes, 2003; Biandolino
and Prato, 2006; Langer et al., 2008) and have reported
the lipid contents to vary between the range of 0.1 to
2.01% in crustaceans. Thus in having lipid content
ranging between 0.23±0.11 to 3.83±0.11%, in muscles the
M. dayanum matches well with other crustaceans. The
lipid content being lower in crustaceans as compared to
molluscs or fin fish, therefore places this organisms in a
better quality food list for human beings.
It could be generalized that fresh water prawns have
higher lipid content than that of marine prawns from the
findings of Chanmugam et al. (1983) wherein total lipid
content of fresh water prawn, M. rosenbergii was found
to be greater than that of marine shrimp i.e., 3.18 to
1.33%. Maximum lipid content in M. dayanum is 3.83%
(Table 1) Perusal of Table 1 showed a rising trend of
lipids in spring (March-April) in ovaries. In extreme
summers (June-July) and winters (Dec-Jan) lipid content
declines to a certain extent. Perusal of Table 1 reveals that
muscle lipid content remains high during November to
January (winter season) and June-July, the period when
ovaries are in quiescent phase. But in concuss with setting
in of ovarian activity the muscle tissue looses lipid and
the ovaries make a matching again (Table 1). Lipids in
muscles are characteristically low in summers. Slight
increase in the lipid content is noticed in spring (April
month) and post monsoon season (Aug-Sept) when they
undergo breeding. Hepatopancreatic lipids showed
characteristically increasing trend in winters with a
decline in spring and summer.
Overall, results indicated that lipid content tends to
increase in muscles and ovary i.e., in spring season
(March-May). Moreover, this season is also their prime
Fig. 3: Showing the stage III (Vitellogenic oocytes) of
reproductive cycle of Macrobrachium dayanum
Fig. 4: Showing the stage IV (Mature oocytes) of reproductive
cycle of Macrobrachium dayanum
concentrically. Nutritive cells arrange themselves along
the periphery of these oocytes but distintive follicular
layer (of nutritive cells) is seen in advanced stages. As the
development advances this layer becomes prominent. Previtellogenic oocytes are seen in different month’s viz.
May to July and December - January.
Stage III: Vitellogenic oocytes (Fig. 3): The oocytes of
this stage are in the range of 230-400 :. The cytoplasm
though vacuolated exhibits granular nature. Small sized
yolk granules make their appearance and spread
centrifugally. There is considerable increase in the size of
the nucleus and the cell. Nuclear membrane becomes
indistinct. Nutritive cells form a distinct epithelium
around the developing oocytes. Maximum numbers of
stage III oocytes are mostly present in June-July and
February-March.
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sites to the gonads for the build up of gametes in
other crustacean species like Penaeus kerathurus
(Mourente and Rodriguez, 1991), Macrobrachium
rosenbergii (Lee and Chang, 1997) Penaeus vannamei
(Palacios et al., 2000), Litopenaeus vannamei
(Wouters et al., 2001), Macrobrachium rosenbergii
(Cavalli et al., 2001), Aristeus antennatus (Rosa and
Nunes, 2003), M. idella idella (Dinakaran and
Sundarapandian, 2009).
breeding season is which energy reserves are required for
gonadal activities.
Monsoon (July to Sept.) is the second short breeding
season of M. dayanum. In this period, muscle and
hepatopancreatic lipid content showed comparatively low
values. Ovaries showed increase in lipid content due to
high gonadal activity in these months for second phase of
breeding. Muscles and hepatopancreas showed low lipid
content in these months could be supported by the fact
that for high gonadal activities, high lipid content in
ovaries are required. This nutrient demand is furnished by
the leaching out of these nutrients from muscles and
hepatopancreas to ovaries.
The observation on the oocytes of M. dayanum
coincides with that of M. kistensis, P. stylifera and
P. hardwickii (Joshi, 1980) and M. lamarrei
(Garimella, 1982). Vitellogenic oocytes have
characteristic feature of having a layer of follicular cells
around the oocyte. Many workers (Joshi, 1980; Garimella,
1982) in the past had also made similar observation.
While investigating the same species, Kailoo (1984) and
Malik (2006) has also reported similar ovarian stages in
M. dayanum. Support to the view point that large amount
of lipids accumulates in muscles during ovarian
quiescence and the same get mobilized from muscle tissue
or other storage organs to ovaries when they are maturing
to cope with reproductive efforts have been advocated in
Common prawn, Palaemon serratus and shrimp, Penaeus
serratus (Soriguer et al., 1997). The Sharp increase in
ovarian lipid accompanying active process of
vitellogenesis thus seems to be closely linked with the
increased rate of fat synthesis in ovaries which demand
and hence mobilize precursor from other generating
and storage tissues of the body. Dramatic increase in
ovarian lipid with maturation could also be justified
by the findings of Lee and Walker (1995) and
Ravid et al. (1999). An important point worth mentioning
here is that hepatopancreatic lipid content is minimum
when stage III and IV oocytes predominates in ovary i.e.
in March-April and August-September and little increases
after spawning (Table 1) suggesting its role in oogenesis
during later stages of the reproductive cycle. These
observations get support from the findings of Teshima and
Kanazawa (1983) who reported that increase of fatty acids
in the ovary has a direct relationship with the decrease of
fatty acids in the hepatopancreas in decapod crustaceans.
Present investigation also showed significant inverse
correlation between muscle lipid and ovarian lipid content
(r = -0.672, p<0.05) throughout the cycle whereas
hepatopancreatic lipid content and ovarian lipid content (r
= -0.186) showed insignificant inverse correlation which
suggests transfer of this fuel to gonads mainly from
muscles throughout the maturation process and from
hepatopancreas in later stages. Several authors have
documented mobilization of lipid reserves from storage
CONCLUSION
Such cyclic variations in lipids recorded during
present studies seen to correspond with seasonal ovarian
stages. Vitellogenic stages of oocytes are prevalent
maximum in the months of March-April and JulySeptember. Accordingly these months are characterized
by a progressive loss in muscle lipid content and a
corresponding increase in gonadal lipid content to be
utilized for vitellogenesis.It is also clear from the Table 1
that ovaries contain higher levels of lipid followed by
hepatopancreas and muscles which suggests that lipids are
important for maturation of crustacean ovaries.
ACKNOWLEDGMENT
The authors are thankful to the Head Department of
Zoology, University of Jammu, for providing the
necessary laboratory facilities.
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AUTHOR’S CONTRIBUTION
Anshu Samyal did sampling and microtomy of
gonads, Yahya Bakhtiyar did the biochemical analysis of
the samples and visited the field, Archana Verma did the
statistical analysis and microphotography, Seema Langer
Supervisor/Guide of this study.
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