Biology of Prawn (Macrobrachium rosenbergii)

All the freshwater prawns that have been cultured so far belong to the genus Macrobrachium , Bate 1868, the largest genus of the family Palaemonidae. About 200 species have been described, almost all of which live in freshwater at least for part of their life.
The giant river prawn, Macrobrachium rosenbergii , was one of the first species to become scientifically known , the first recognizable illustration appearing in 1705. The nomenclature of freshwater prawns, both on a generic and a species level has had quite a muddled history. In the past, generic names have included Cancer (Astacus) and Palaemon. Previous names of M. rosenbergii have included Palaemon carcinus, P. dacqueti and P. rosenbergii ( De Man 1879) became universally accepted.

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Biology of Prawn (Macrobrachium rosenbergii)

Introduction:
All the freshwater prawns that have been cultured so far belong to the genus Macrobrachium ,
Bate 1868, the largest genus of the family Palaemonidae. About 200 species have been
described, almost all of which live in freshwater at least for part of their life.
The giant river prawn, Macrobrachium rosenbergii , was one of the first species to become
scientifically known , the first recognizable illustration appearing in 1705. The nomenclature of
freshwater prawns, both on a generic and a species level has had quite a muddled history. In the
past, generic names have included Cancer (Astacus) and Palaemon. Previous names of M.
rosenbergii have included Palaemon carcinus, P. dacqueti and P. rosenbergii ( De Man 1879)
became universally accepted.
There are 200 species of Macrobrachium in the world, of which 49 are commercial. Twenty
seven of the commercial species are found in Asia and the Pacific. Most live in freshwater. A
few species live in saltwater in the mouths of rivers.
M. rosenbergii is found extensively in the tropical and subtropical waters of the Indo-Pacific
region in Malaysia, Thailand, the Philippines, India, Shri Lanka, Bangladesh, Myanmar,
Indonesia and Vietnam. They are generally found in freshwater, in ponds, rivers, lakes, ditches,
canals, depressions, low-lying floodplains and river mouths. Most of the species spend their early
life in saltwater that is connected directly or indirectly with the sea. Some species complete their
life cycle in freshwater, but these are not of commercial importance.
Prawns move upstream, entering lakes and even paddy fields, up to about 200 km from the sea.
This type of migration is observed not only in M. rosenbergii but also in other species of
Macrobrachium.

Importance:
Prawns are important groups of shellfish belongs to the class crustacean. Prawns have large
economic significance. So, it is cultured worldwide. In Bangladesh, there are 24 species of
freshwater prawns. It is very important to know about their external and internal morphology. On
the other hand, male and female prawn have some distinct external characteristics between them.
-To know their external and internal morphologies
-To know about their maturity and their reproduction techniques
-To distinguish male and female
-To determine their culture techniques
-To know about their feeding and breeding behavior
Subspecies of M. rosenbergii

Due to differences in climate, weather and natural environment, many subspecies of M.
rosenbergii have evolved. Three varieties are generally observed in nature.

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Blue claw subspecies:
This subspecies grows to a large size. The ratio of claw to body length is 1 .6 ±0. 1. The male
is territorial, its breeding behavior is complex and growth is comparatively slow.

Orange claw subspecies:
This subspecies is a little bigger than the median size of the blue claw variety and has orange-
colored claws. The ratio of claw to body length is 1.0 ±0.05. The rate of fertilization of eggs is
comparatively slow, but growth is fast.

Small subspecies:
This is the smallest of all the subspecies with spineless claws. The ratio of claw to body length
is 0.5 ±0.1. At the time of copulation, these adopt the ‘snake’ mating strategy. Their growth is
the slowest of the three varieties.

Classification:
Kingdom: Animalia
Phylum: Arthropoda
Sub-Phylum: Crustacea
Class: Malacostraca
Order: Decapoda
Family: Palaemonidae
Genus: Macrobrachium
Species: M. rosenbergii


Description:
The body of post larval and adult prawns consists of the cephalothorax (head) and the abdomen
(tail). The bodies of freshwater prawns are divided into twenty segments (known as somites).
There are 14 segments in the head, which are fused together and invisible under a large dorsal
and lateral shield, known as the carapace. The carapace is hard and smooth, except for two
spines on either side; one (the antennal spine) is just below the orbit and the other (the hepatic
spine) is lower down and behind the antennal spine. The carapace ends at the front in a long beak
or rostrum, which is slender and curved upwards. The rostrum extends further forward than the
antennal scale and has 11-14 teeth on the top and 8-10 underneath. The first two of the dorsal
teeth appear behind the eye socket.
The front portion of the cephalothorax, known as the cephalon, has six segments and includes the
eyes and five pairs of appendages. The final three of these six segments can be seen if the animal

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is turned upside down and appendages of the thorax are moved aside. The cephalon segments
therefore support, from the front of the animal.
 The stalked eyes;
 The first antennae, which each have three-segment peduncles from which three tactile
flagella emerge;
 The second antennae, which each have five-segments peduncles and a single long
flagellum;



Figure 1.1: External morphology of freshwater prawn M. rosenbergii


 The mandibles, which are short and hard and are used to grind food;
 The first maxillae, which are plate-like, hidden below the second maxillae, and used to
transfer food into the mouth; and
 The second maxillae, which are similar to the first maxillae but have an additional
fuction.
The rear portion of the cephalothorax, known as the thorax, consists of 8 fused segments which
have easily visible pairs of appendages. These appendages consist of 3 sets of maxillipeds and 5
pairs of pereiopods, as follows:

 The first and second maxillipeds are similar to the first and second maxillae and function
as mouthparts;
 The third maxillipeds, which are also mouthparts but look rather like legs;
 The first and second legs, which have pincers. These pincerened legs are also called
chelipeds.

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 The second chelipeds are used for capturing food, as well as in mating and agonistic
behavior ; and the third, fourth and fifth legs (pereiopods), which are much shorter than
the second cheliped, have simple claws, and are sometimes called walking legs. Eggs are
extruded from oval gonophores in the base of the third pereiopods of females, which are
covered with a membrane. In males, sperm is extruded from gonophores which are
covered by flaps, situated in the base of the fifth pereiopods.





M. rosenbergii can grow to a length of over 30 centimeters (12 inch). M. rosenbergii has been
used in research more than any other species and has been introduced many new countries for
commercial culture. Fujimura and Okamoto (1972) were successful in producing post-larvae
(FL) of M. rosenbergii in large numbers in Hawaii in 1972. M. rosenbergii is being cultured in
commercial quantities in many parts of the world including Hawaii, Honduras, Mauritius,
Taiwan, Thailand and the Philippines. Farms have also been developed in Costa Rica, Israel,
Bangladesh, Malaysia, and Mexico.

The shape (external morphology) and other characteristics of M. rosenbergii:
Eggs of M. rosenbergii are slightly elliptical, with a long axis of 0.6-0.7 mm and are bright
orange in color until 2-3 days before hatching when they become grey-black. This color change
occurs as the embryos utilize their food reserves. Most scientists accept that the larvae go
through 11 distinct stages (Uno and kwon 1969) before metamorphosis, each with several
distinguishing features which are described and illustrated in Table 1.


Figure 1.2: Grilled giant river prawns in Thai cuisine; each
(whole) prawn weighing around 500 grams

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Selected characteristics of M. rosenbergii:


Table 1

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M. rosenbergii go through eleven distinct larval stages (Figures 1-11) before metamorphosing to
become post larvae:

Figure 1.3: Eleven distinct larval stages before metamorphosing to become post larvae M.
rosenbergii

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A summary of the segments and the functions of each appendages is provided in; Table 2 below:

Body section Somites Appendage Name(Pair) Function of appendages and related
structures



Cephalon:
Front portion of the
cephalothorax
1
Embryonic segment

2 1
st
antennae

Tactile & sensory perception
3 2
nd
antennae

(statocyst)
4 mandibles

Tactile
5 1
st
maxillae

Cutting & grinding food
6 2
nd
maxillae

Food handling; water circulation through
gill chamber





Thorax:
Rear portion of the
cephalothorax
7 1
st
maxillipeds

Feeding/ food handling
8 2
nd
maxillipeds

Feeding/ food handling
9 3
rd
maxillipeds

Feeding/ food handling
10 1
st
pereiopods

Food capture
11 2
nd
pereiopods

Food capture; agonistic and mating
behavior
12 3
rd
pereiopods

Walking; female gonophores between
base of legs
13 4
th
pereiopods

Walking
14 5
th
pereiopods

Walking; male gonophores between base
of legs




Abdomen
15 1
st
pleopods

Swimming
16 2
nd
pleopods

Swimming; copulation in males: appendix
masculine
17 3
rd
pleopods

Swimming
18 4
th
pleopods

Swimming

19
5
th
pleopods Swimming

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Abdomen

20

Uropods

Propulsion; together with the central
telson.

Identifying M. rosenbergii from other freshwater prawn species:
 It has a very long rostrum, with 11-14 dorsal teeth and 8-10 ventral teeth.
 The tip of its telson reaches distinctly beyond the posterior spines of the telson;
 The adult male has very long second chelipeds in which all segments are elongate and
have blunt spines;
 The movable finger of the second chelipeds of the adult male is covered by a dense
velvet-like fur but this fur is absent from the fixed finger and the rest of the cheliped; and
 It is the largest known of all Macrobrachium species, adult males having been reported
with a total body length of up to 33 cm, and adult females of up to 29 cm.

The internal morphology and other characteristics of freshwater prawn M. rosenbergii:

Excretory system:

 Several tissues and organs participate in the excretion of metabolic wastes in crustaceans,
but the primary organ of urine production in adult malacostracans is the antennal gland/
green gland, which is paired and located in the cephalothorax.
 The morphology of this gland follows a basic plan: each consists of an initial sac, a long
excretory tubule, and a large bladder. The opening is an excretory pore near the base of
the antennae.
 Urine is formed in the antennal gland by filtration and reabsorption, with a tubular
secretion added. Although the antennal gland is dominated as the excretory organ, the site
of nitrogenous waste excretion is actually the gills.
 M. rosenbergii 32
0
C as ammonia-N, 25.6% as organic-N and 4.2% as urea-N.


Reproductive system:

 The internal reproductive structures of M. rosenbergii are located in the cephalothorax.
 In males, they consist primarily a pair of testes, which are fused and lie mid dorsally in
the cephalothorax, giving rise to vas deferens. The paired vasa deferentia are simple tubes
that end in terminal ampule, which contain the spermatophores and open at the
gonophores on the coxae of the 5
th
pereiopods. During mating, the ampullae extrude the
spermatophores containing the sperm mass.
 Besides these structures, an androgenic gland is attached to the vasa differentia ; this is
involved with the development and regeneration of secondary male sexual characteristics
and also controls morphotypic differentiation.

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 In females, the paired ovaries are located dorsally to the stomach and hepatopancreas.
They give rise to a pair of oviducts which extends towards and open into, the gonophores
on the basal segment of the 3
rd
pereiopods.


Reproduction:
Sexual dimorphism
M. rosenbergii can sexually distinguished with first appearance of gonophores in juveniles, at
5.9 mm for males and 7.6 mm for females. Among M.
rosenbergii’s five pairs of walking legs, the second is the
biggest and has a pincher (chela) at its tip. Both legs of
this second pair are of the same size. Mature M.
rosenbergii males are bigger than the females, with their
cephalothorax larger and their second pair of thoracic legs
comparatively longer and thicker. The cephalothorax of
the male is also proportionately larger and the abdomen
narrower than the female’s. The genital pores of the male
are situated at the base of the fifth pair of walking legs. In
immature males, there is a raised hard point on the first
segment of the abdominal part of the body. Genital pores
of the female are situated at the base of the second pair of
thoracic legs. The abdominal pleura of the Female are
comparatively longer and the abdomen wider. The orange-
colored maturing gonad is easily visible. It is relatively
easy to differentiate between deheaded freshwater prawn
and marine shrimp. In freshwater prawn, the second
abdominal pleuron overlaps the first and third pleura. In
marine shrimp, the second pleuron overlaps only the third
pleuron and is itself overlapped .


Reproductive behavior:
After ovarian maturation, M. rosenbergii females experience a moult, known as a pre-spawning
or pre-mating moult, which usually occurs at night. After this moulting process, courting and
mating commence. Successful mating can only take place between ripe females, which have just
completed their pre-mating moult and are therefore soft-sheltered, and hard-shelled males.
Mating of M. rosenbergii is completed in six stages, which are summarized below:
(1) The female approaches a BC male.
(2) The female climbs over the telson of the male. Antennal contact is maintained at all times
by sweeping between and female at this stage.
Figure 1.4: Identifying characteristics of male
& female

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(3) The female orientates herself to the front of the male and moves backwards until she is
between his chelae.
(4) Once between his chelae, the female may turn to face the male, making contact with the
male’s walking legs. Prior to fertilization, the female re-orients herself in order to
position her dorsal telson under his head region.
(5) The male mounts the female, and begins characteristics rubbing action with pleopods on
the ventral lower cephalothorax and upper abdominal segments of the female. The female
appears to become torpid.
(6) The male grasps the female’s rostrum with his chelae of pereiopods, and may also grasp
the gill operculum. The female is turned ventral side up with chelae stretched out in front,
and remains torpid. This ensures that the reproductive organs are aligned.

Life history:
There are four stages in the life of a freshwater prawn, viz, egg, larva, juvenile and adult
(See Figure 1.5). Like other crustaceans, the freshwater prawn moults. The number of moults and
the durations of intermoults are not fixed, and depend on the environment, particularly
temperature and the availability of food.
In the natural environment, mating of Macrobrachium takes place all year round, although, due
to environmental reasons, peak mating takes place only during certain periods of the year. A
female prawn, with matured gonad, copulates just after moulting with a male prawn having a
hard shell. During copulation, the male deposits a gelatinous mass, or spermataphore, on the
underside of the thorax of the female, between her walking legs. The female prawn releases its
eggs a few hours to a few days after copulation. The number of eggs depends on the size of the
female. A fully matured female of 50-100 g can carry 50,000-100,000 eggs. But at first maturity,
due to the female’s small size, it lays only 5000-20,000 eggs.
As the eggs are extruded from the gonophore, they are fertilized by non-motile sperm retained
in the spermatophore. The fertilized eggs are then transferred to a brood chamber on the
underside of the abdominal region of the female, held in place by a thin membrane and kept
aerated by vigorous movement of the abdominal appendages. Eggs are incubated in this way for
21 days and then hatch. In the laboratory, it has been observed that hatching takes place 20 days
after copulation; it may even take 25-30 days if the temperature has remained below 28°C.
Ovaries frequently ripen again even while a female is carrying eggs. Immediately after hatching,
the female can again release these eggs. In some cases, a female can lay eggs twice a month. The
eggs of the prawn are slightly elliptical, the longer axis being 0.6 - 0.7 mm in length. They are
bright orange in color until two or three days before hatching, when they become slate gray.
Larvae hatch during the night. Rapid movement of the female pleopods disperses the newly
hatched larvae, which normally swim with their heads down and ‘jump’ when they contact a
surface. Larvae need brackish water to survive at this stage. Even if larvae hatch in freshwater,
they will not survive if they are not put into brackish water within two or three days. Larvae in
the wild generally eat zooplankton, small insects and larvae of other aquatic invertebrates.
Larvae in a hatchery take a minimum of 26 days to metamorphose into post-larvae (PL). Post
larvae can tolerate a wide range of salinity, but freshwater is their normal habitat. And so, two to
three weeks after metamorphosis, the PL move against the current and head towards freshwater
canals and rivers. They abandon the planktonic habit at this stage and become omnivorous,

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feeding on aquatic insects and their larvae, phytoplankton, seeds of cereals, fruit, small mollusca
and crustacea, fish flesh, slaughterhouse waste and animal remains. They move by crawling and
generally swim with their dorsal side uppermost. They can swim rapidly.


Figure 1.5: The life cycle of M. rosenbergii


Food and feeding habit:

M. rosenbergii are omnivores and mostly nocturnal feeder. As a larvae grow they become
aggressive sight feeders and feed almost continuously, primarily on small zooplankton, worms
and larval stages of other aquatic invertebrates.
In addition to the types of food they ate as larvae, post larval prawns eat larger pieces of animal
and plant materials will be ingested. The diet includes larval and adult insects, algae, mollusks,
worms, fish and faces of fish and other animals.
At high densities, or under conditions of food limitations, prawns become cannibalistic.


Moulting
Moult Cycle:
Although moulting appears to take place as a single event, it is a multistage process. The moult
cycle in M. rosenbergii has been defined as:
i. Post- moult,
ii. Inter-moult,
iii. Pre-moult and
iv. Moult.

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These stages might not be distinguished very clearly. But the behavioural differences during
different stages can easily be observed.
The general physiological changes during different stages in a moult cycle are given below:
POST-MOULT (METAECDYSIS/SOFT BACK STAGE)
This stage falls a few days after moulting. This stage is characterized by hardening of the newly
formed exoskeleton by deposition of calcium and other minerals and results in the complete
formation of a new exoskeleton.

Figure 1.6(a) Entire shed-off shell of a prawn



INTER-MOULTING (ANECDYSIS)
This is the stage between successive moults. The prawn in this stage is called an instar. The
water taken up during ecdysis is gradually replaced by tissue growth, enrichment of organic
matter and mineral reserves, resulting in growth of the prawn.

Figure1.6 (b) Male morphotypes of prawn

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PRE-MOULT (PROECDYSIS/ PEELER STAGE)
This is the stage just before moulting. In this stage, calcium is reabsorbed from the old
integument leading to enlarged epidermal cells. There will be clear separation of the old
integument from the epidermis for moulting. Appendages also regenerate.
MOULT (ECDYSIS)
The animal comes out of the old exoskeleton with a newly formed soft exoskeleton underneath
and absorbs water in the tissue for increase in size. The above stages are broad classification of
the moult cycle, which give some idea of the physiological process that takes place during
moulting.

In M. rosenbergii, the length of the moult cycle varies depending on the size, sex and age of the
animal. However, it is not clear about the exact duration of each stage. In Penaeids generally the
moult cycle is short for juveniles and long for adults. Giant freshwater prawn exhibits long
(about 30-80 days) inter-moult periods, short (about 10-12 days) pre-moult and very short (about
2-6 days) post-moult periods. In Penaeus merguiensis, actual moulting lasted for 40 seconds and
in P. duorarum it is only 20-30 seconds.


How to distinguish male and female of M. rosenbergii ?
The difference between male and female species are given below:

Male Female
a. Body larger than female a. Body usually smaller than male
b. 2
nd
walking leg is much larger b. 2
nd
walking leg is short & thin
c. It has appendix masculine in pleopod c. Brood chamber in pleopod
d. Cephalothorax is larger than female d. Cephalothorax is smaller
e. In juvenile stage, 1
st
appearance of
gonophore at 5.9 cm
e. 1
st
appearance of gonophore at 7.6 cm
f. 1
st
abdominal segment has pointy
central lump
f. Female has no central lump
g. Adult male is bluish-dark in color g. Female abdomen is in orange color
during breeding season


How to distinguish Macrobrachium and penaeid (marine) shrimp ?

Many people find it hard to distinguish between Macrobrachium and penaeid (marine) shrimp,
once they have been harvested and the heads have been removed. If the ‘tail’ still retains its shell
there are, in fact, two easy ways of distinguishing them (Fincham and Wickins, 1976).

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Firstly, Macrobrachium spp, have a smooth rounded dorsal surface to the abdomen while
penaeids have a simple or complex ridge at the dorsal apex of the abdomen. (Figure 1.7)


Figure 1.7

Secondly, the second pleuron of the abdomen of Macrobrachium overlaps both the first and the
third pleuron. In penaeids the second pleuron overlaps the third pleuron only and is itself
overlapped by the first. (Figure 1.8)



Figure 1.8


Recommendation:

There is endless possibilities to make this fish farming sector very economically important.
Country like Bangladesh is agriculture based country where having lots of opportunity to
develop a well infrastructure of shellfish farming industry. We are earning a lot of foreign
exchange by exporting M. rosenbergii to Europe, America and other countries of the world.
Some workshops could be done from field level farmers to give them a clear idea about shellfish
farming. Government and NGO’s are important factors here because they can keep contribution

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from field level. M. rosenbergii has been used in research more than any other species and has
been introduced many new countries for commercial culture. Fujimura and Okamoto (1972)
were successful in producing post-larvae (FL) of M. rosenbergii in large numbers in Hawaii in
1972. M. rosenbergii is being cultured in commercial quantities in many parts of the world,
including Hawaii, Honduras. Mauritius, Taiwan, Thailand and the Philippines. Farms have also
been developed in Costa Rica, Israel, Malaysia, and Mexico.

Conclusion:
It is necessary to have clear knowledge to culture prawns without clear knowledge no person can
do it successfully. As Bangladesh is a developing country, our hatchery operators should try to
enhance the prawn production. We can earn more foreign exchange by doing culture prawn
intensively. Every fisheries related person should come out to help and contribute in this
increasing sector.

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