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PENAEUS MONODON
Penaeusis a genus of prawn under the class Crustacea. It has a worldwide distribution.
The various species belonging to this genus are found both in tropical and temperate latitudes.
Practically all of them are marine although some are known to spend a part of their life in the
brackish water and even in freshwater.Some species represented in Indian waters are Penaeus
japonicus, P.latisulcatus, P. monodon, P.semisulcatus, P. indicus, P. merguiensls and P.
perucillatus.
External characters; The body of. Penaeus consists of two regions, the anterior
cephalothorax and the posterior abdomen
Cephalothorax:
1. The cephalothorax is formed by the fusion of the head or cephalon and thorax.
2. The head consists of five segments and the thorax is eight-segmented.
3. The dorsal and the lateral sides of the cephalolhorax is covered by- a single
chitinous exoskeleton called carapace or dorsal shield. It is formed by the fusion
of five terga of the head and eight terga of the thorax.
4. A transverse cervical groove on the dorsal side seperates the anterior head from
the posterior thorax.
5. The anterior region of the carapace forms in front a median serrated process
known as rostrum. There are two compound eyes, which are attached to the base
of the rostrum by movable stalks.
6. The free ventral flaps of the carapace on each side of the thorax is called.
branchiostegite or gill cover.The space between the branchiostegite and body
wall of each side is known as branchial chamber.
7. The respiratory organ, branchiae or gills lie in the branchial chamber.The ventral
sterna of the cephalothorax fuse and form a plate known as sternal plate.
8. In female, the sternum of the last thoracic segment forms an outgrowth called
thelycum,which encloses acavity.The male deposits its spermatophores into it.
Cephalothorax bears., thirteen pairs of jointed appendages, of which the anterior
five pairs are in the the cephalic region and the posterior eight pairs are in the
thoracic region.
9. Infemale,a pair of genital openings lies at the base,of the sixth thoracic legs. In
male, a pair of genital aperturelies on a small papillae at the base of last pair of
thoracic legs.
Abdomen:
1. The abdomen consists of six segments and a terminal piece called telson.
2. The dorsal side of each abdominal segment is covered by an exoskeleton called tergum .
Appendages:
Cephalic Appendages:
Antennules:They are the anterior most appendages situated below the level of the
eyestalks. They are also known as first antennae. Each antennule has a proterpodite, an exopodite
and an endopodite .Protopodite has three podomeres, the proximal precoxa, middle coxa and the
distal basis. The precoxa has a hollow depression on one side. The eye and the balancing sense
organ/ statocyst are located in it. The opening of the statocyst lies at the base of this podomere.
The basis carries short many jointed flagella-like exopodite and endopodite. They function as
tactile sense organs.
Antennae:They are also known as second antennae. They lie immediately behind the
antennules. Each antennule consists of protopodite, exopodite and endopodite. Protopodite has
two podomeres, the proximal coxa and distal basis. The opening of the excretory duct lies in the
coxa. Exopodite and endopodite are situated on the basis. Endopodite has three basal podomeres
and a long narrow many-jointed filament. It functions as tactile sense organ. The exopodite is a
flat broad plate-like structure and it is also known as squame. It functions as a balancing organ.
Mandibles:They lie on either side of the mouth. Each mandible has a protopodite and an
endopodite. Exopodite is absent. Protopodite is a single stout calcified structure with toothed
inner edge.The teeth are used for grinding food to size. The endopodite, which is attached to the
protopodite is segmented. The endopodite is also known as sensory palp.
First pair maxillae (Maxillulae):They are the smallest appendages of prawn. Each
maxilla has a protopodite and an exopodite.Exopodite is absent. Each protopodite has two
flattened leaf-like lobes, the coxa and basis. They protect inwards as jaws and are also called
gnathobases.Endopodite is uniointed and leaf-like with narrow distal end. Both gnathobases and
endopodite are provided with sharp hair like processes.
Second Pair maxillae:They lie just behind the first Pair of maxillae. Eacir maxilla is
provided with a protopodite, an exoPodite and an endopodite.
Protopodite is flat and four lobed.Their edges are flattened and provided with hair-like
recesses.They aid in mastication.The endopodite is small, unsegmented and leaf-iike. It is
situated between the protopodite and the exopodite.The exopodite is large, expanded, flat and
boat-shaped and is also known as scapho gnathite. Its movement produces. Water currrents in
the gill chamber.
Peraeopods (walking legs):There are five pairs of peraeopods. The first three pairs of
peraeopods are known as chelate legs or chelipeds and the last two pairs of peraeopods are
known as non-chelate legs. In all peraeopods, the exopodite is small and fringed with many hair-
like structures.
Chelate legs (Chelipeds):All chelate legs are identical in structure. But they differ
slightly in size, the third one being the longest. The five jointed endopodite shows chelate
articulation of the terminal two podomeres, the propodus and the dactylus . By the hinged
articulation of the dactylus to the side of the propodus, a pincer like apparatus is formed. This
helps in grasping the food and passing it on to the mouth. The chelate legs are also used for
walking
Non-chelate legs;In non-chelate legs, dactylus and propodus do not form chelate
articulation . Epipodites are absent. Non-chelate legs are used for walking.
1. First pleopods:In female, exopodite is flattened, thick and fringed with setose processes.
Endopodite is absent or may be present as a very small bud-like process. In male,
endopodite is short and provided with hooks.The hooks of the endopodites of the two
sides interlock and form a rod-like structure called the petasma which is used for
transferring sperms into the thelycum of female.
2. Second, to fifth pleopods:The second, third, fourth and fifth pairs of pleopods have a
typical biramous structure, with a two-jointed protopodite, and unjointed exopodite and
endopodite .
The digestive system of Penaeus consists of the alimentary canal and the digestive gland .
Alimentary canal;The alimentary canal is a long tube, which extends from the mouth to
the anus. It is divisible into three regions, stomodaeum, mesenteron and proctodaeum. The
stomodaeum is the anterior region and it is lined with chitin. It consists of the mouth, the buccal
cavity, the oesophagus and the stomach. The mouth lies on the ventral side of the cephalothorax,
between the mandibles. It leads into a short buccal cavity. The stomach and the buccal cavity are
connected by a short vertical tube called the oesophagus. The stomach consists of an anterior
larger cardiac stomach and the posterior smaller pyloric stomach. The inner ventral folds of the
cardiac stomach are provided with setae and spicules. The inner dorso-lateral folds of this region
bear stout denticles which form the gastric armature or gastric mill for grinding the food. The
pyloric stomach encloses a narrow cavity.
The pyloric stomach opens into the mesenteron.The mesenteron is the middle region, of
the alimentary canal. It is not lined with chitin.The chitinous lining of the pyloric stomach forms
one median dorsal,one median ventral and two lateral lippets or valvulae.They project into the
mesenteron.The proctodaeum is the posterior region of the alimentary canal.It is lined
with chitin. It consists of the hindgut (or rectum) and the anus.The anus lies at the base of
the telson ventrally.
Circulatory system
Circulatory system consists of heart,arteries,pericardial membrane, pericardial
sinus,haemocoel , bood channels and blood or haemolymph. The heart is a triangular chamber. It
lies in the pericardial space.,It is provided with paired openings called ostia. Arteries are the
main tubes which arise from the anterior and posterior regions of the heart. The pericardial
membrane lies below the dorsal body wall just above the alimentary canal. The space between
the dorsal body wall and the pericardial membrane is the pericardial sinus.The spaces between
the visceral organs form the haemocoel. It contains blood or haemolymph. From the haemocoel
blood goes to the gills through the blood channels. From the gills blood goes to the heart through
blood channels. The blood contains plasma, haemocytes or blood cells and the respiratory
pigment haemocyanin.
Circulation of haemolymph
Respiratory system
Penaeus takes up oxygen dissolved in sea-water. Its respiratory organs are inner lining of
branchiostegites, epipodites (mastigobranchiae) and branchiae (gills) .
Branchiae or Gills:Branchiae (gills) are the feather like (plumose ) outgrowth of the
lateral wall of the thorax and thoracic appendages. Each gill has a stem Three longitudinal blood
channels run through the stem. They are two lateral channels and one median channel. The two
Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016
lateral channels are connected by many transverse channels. Many lateral flat gill plates arise
from the stem.Marginal channels from the lateral channels penetrate into the gill plate and open
into the median channel, The stem and gill plate are covered externally a thin layer of chitin. A
single layer of epithelial cells lies beneath it. Epithelial layer encloses connective tissue and
blood channels. This kind of gill is known as dendrobranchia.
According to the point of origin, there are three sets of branchiae. They are:
Branchial Formula:The formula, which shows the number and arrangement of the
branchiae or gills on one side of the thorax is known as branchial formula.
Mechanism of Respiration
There are two branchial or gill chambers. They enclose highly vascular respiratory organs
(branchiae or gills, epipodites and inner lining of the brachiostegites). The anterior,ventral and
posterior sides of each chamber are open. The exopodite (scaphognathite) of the second maxilla
lies at the anteriorregion of the bianchial chamber. Its movement drives water out of the
branchial chamber. So water from outside enters the chamber through the posterior side.A
constant current of water flows over the respiratory organs.Oxygen dissolved in the water
diffuses into the blood in the respiratory organs and carbon dioxide in the blood diffuses into the
water. The setose processes along the anterior, ventral and posterior margins of the branchial or
gill chambers prevent the entry of foreign particles into the chambers.
Sense organs;The receptor organs of Penaeus are compound eyes, statocysts, tactile organs and
setae.
Compound eyes:There are two compound eyes/ one on either side of the base of the rostrum.
They are at the ends of movable stalks. Each eye is made up of many optical units called simple
eyes or ommatidia. The ommatidia are elongate, rod- shaped and radially placed. They lie side-
by-side and are separated from each other by a sheath of black pigment cells. Each ommatidium
consists of an outermost layer called cornea. It is formed by the transparent cuticle. Externally
this layer is hexagonal. Each hexagonal part is called a facet. Corneagen layer lies below the
cornea.It consists of two epidermal cells or lenticular cells. They secrete the outer cornea. Four
cone cells lie beneath the corneagen layer and they are known as vitrellae or crystalline cone
cells. The central transparent regions of. the four crystalline cone cells form the crystalline
cone.The crystalline cone cells of the ommatidium are surrounded by distal pigment cells which
contain dark brown or black screening pigment. Cornea, corneagen layer, cone cells, crystalline
cone and distal pigment sheath form the focussing or dioptric apparatus of the
ommatidium.Beneath the cone cells lie seven elongated cells retinular cells .They form the
retinula. The microvilli arising from the axial surface of the retinular cell interdigitate with those
of adjacent retinular cell and form a rhabdome in the central region of the retinula. The retinular
cells contain proximal screening pigment. Retinular cells and rhabdome form the receptive or
retinal part of the ommatidium.All the ommatidia rest on a porous basal lamina. From the basai
region of each retinular cell arises an axon or nerve fibre. It passes through the pore. The axons
of all the ommatidia of an eye form the optic nerve. Two types of images are formed by the
compound eye of Penaeus.They are mosaic (apposition) and super-imposed (superposition)
images.
Mosaic (apposition) image:Each ommatidium receives light rays from a small part of an object
direct in front of it. Oblique rays are absorbed by the pigments. Straight light rays which are
parallel to the axis of the ommatidium reach the rhabdome and form image of that part. The
images of different parts of the object, formed by the different ommatidia together form the
mosaic or apposition image of the object.This image is erect and not inverted.
Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016
Super-imposed (superposition) image:In dim light, the pigments in the distal pigment cells and
the retinular cells are concentrated in their respective cells. Thus the optical separation between
adjacent ommatidia is removed. Each ommatidium receives light rays from an object in front of
it through the facets of the surrounding ommatidia as well as from its own and forms an image.
Images formed by the ommatidum are the result of the superposition of light rays from a number
of facets.The compound image thus formed is known is as superposition image.
Tactile organs and setae :Antennae are the important tactile sense organs. Many sensory setae
are located over the body surface especially the appendages.
Reproductive system:Male and female sexes are separate (gonochoric) and they are sexually
dimorphic. The female can be distinguished from the male by. the following external characters.
All freshwater prawns (like other crustaceans) have to regularly cast their ‘exoskeleton’
or shell in order to grow. This process is referred to as moulting and is accompanied by a sudden
increase in size and weight. There are four distinct phases in the life cycle of the freshwater
prawn, namely eggs, larvae, postlarvae (PL) and adults. The time spent by each species of
The life cycle of M. rosenbergii can be summarized as follows. The mating (copulation)
of adults results in the deposition of a gelatinous mass of semen (referred to as a spermatophore)
on the underside of the thoracic region of the female’s body (between the walking legs).
Successful mating can only take place between ripe females, which have just completed their
pre-mating moult (usually at night) and are therefore soft-shelled, and hard-shelled males. All of
the various types of males are capable of fertilising females but their behaviour is different (see
Annex 8). Detailed descriptions of the mating process are given in Ismael and New (2000) and
Karplus, Malecha and Sagi (2000). Under natural conditions, mating occurs throughout the year,
although there are sometimes peaks of activity related to environmental conditions. In tropical
areas these coincide with the onset of the rainy season, whereas in temperate areas they occur in
the summer.
Within a few hours of copulation, eggs are extruded through the gonopores and guided by
the ovipositing setae (stiff hairs), which are at the base of the walking legs, into the brood
chamber. During this process the eggs are fertilized by the semen attached to the exterior of the
female’s body. The eggs are held in the brood chamber (stuck to the ovigerous setae) and kept
aerated by vigorous movements of the swimmerets. This is in contrast to penaeid shrimp, whose
fertilized eggs are released into the sea, where they hatch. The length of time that the eggs are
carried by female freshwater prawns varies but is not normally longer than three weeks. The
number of eggs which are laid depends also on the size of the female. Female prawns of M.
rosenbergii are reported to lay from 80 000 to 100 000 eggs during one spawning when fully
mature. However, their first broods, (i.e. those which are produced within their first year of life),
are often not more than 5 000 to 20 000. Females normally become mature when they reach 15-
20 g but berried females have been observed as small as 6.5 g (Daniels, Cavalli and Smullen
2000). Under laboratory conditions, where a breeding stock of both males and females was kept,
it has been noted that egg incubation time averaged 20 days at 28°C (range 18-23 days). Ovaries
frequently ripened again while females were carrying eggs. Pre-mate intermoults were separated
by as little as 23 days (i.e. females on some occasions hatched two batches of eggs within a one-
month period). It is unlikely that this would happen under natural conditions but it does show the
potential fecundity of the animal.
As the eggs hatch, a process which is normally completed for the whole brood within one
or two nights, the larvae (free-swimming zoeae) are dispersed by rapid movements of the
abdominal appendages of the parent. Freshwater prawn larvae are planktonic and swim actively
tail first, ventral side uppermost (i.e. upside down). M. rosenbergii larvae require brackishwater
for survival. Those which hatch in freshwater will die unless they reach brackishwater within a
few days. There are a number of microscopically distinct stages during the larval life of
freshwater prawns, which lasts several weeks (Annex 1). Individual larvae of M. rosenbergii
have been observed, in hatchery conditions, to complete their larval life in as little as 16 days but
reaching this stage may take much longer, depending on water temperature and other factors.
The importance of this observation is fully discussed later in the manual. Larvae eat continuously
On completion of their larval life, freshwater prawns metamorphose into postlarvae (PL).
From this point onwards they resemble miniature adult prawns and become mainly crawling
rather than free-swimming animals. When they do swim it is usually in a normal (dorsal side
uppermost) way and in a forward direction. Rapid evasive movement is also achieved by
contracting the abdominal muscles and rapid movement of the tail fan. Postlarvae exhibit good
tolerance to a wide range of salinities, which is a characteristic of freshwater prawns.
Postlarvae begin to migrate upstream into freshwater conditions within one or two weeks
after metamorphosis and are soon able to swim against rapidly flowing currents and to crawl
over the stones at the shallow edges of rivers and in rapids. They can climb vertical surfaces and
cross land, provided there is abundant moisture available. In addition to using the foods available
to them as larvae, they now utilize larger pieces of organic material, both of animal and
vegetable origin. Postlarval freshwater prawns are omnivorous and, as they grow, their natural
diet eventually includes aquatic insects and their larvae, algae, nuts, grain, seeds, fruits, small
molluscs and crustaceans, fish flesh and the offal of fish and other animals. They can also be
cannibalistic. Further reading on this topic may be found in Ling (1969).