Indian Journal of Experimental Biology

Vol. 43, March 2005, pp. 209-223

Review Article

Current progress in shrimp endocrinology-A review

ADDiwan
Indian Council of Agricultural Research (ICAR), Krishi Anusandhan Bhavan-II, Pusa, New Delhi 110 012, India

One problem in aquaculture is obtaining brood because many commercially important species are incapable of
spontaneous maturation under artificial conditions. Commercial shrimp hatcheries commonly use eyestalk ablation to
stimulate gonadal maturation in shrimps. Research has been conducted on the inhibition of reproductive maturation by
hormones originating in the eyestalk glands and on other endocrine sources (e.g.,brain, thoracic ganglion, ovary, mandibular
organ, androgenic gland and Y-organs) to determine their roles. Alternate techniques for acceleration of gonad maturation
through the use of synthetic hormones or neurotransmitters may benefit aquaculture. Neurohormones and neuroregulators
have been shown to accelerate gonadal maturation but an effective delivery technique must be developed for use in a large-
scale aquaculture operation.

Keywords: Shrimp, Hormones, Neurotransmitters, X-organ, Y-organ, Sinus gland

Shrimps are common to worldwide capture fisheries
and to aquaculture. Demand for shrimp in the
international markets has increased greatly over time.
The high demand has led to over-exploitation of the
natural shrimp stocks in many parts of the world. A
decline in the capture fisheries led to increased
aquaculture production as an alternative source for
sustaining and expanding the seafood industry. Due to
overexploitation of the wild stock of marine shrimp
and the decrease in total natural productivity, shrimp
farming activity has developed rapidly in two
decades, with an the annual global production of
700,000 mtl. Shrimp culture has many challenges.
One major problem is the lack of genetic selection of
brood. Zeleny2 and Panouse3 showed the effect of
eyestalk ablation on moult and ovary development in
some crustaceans. Reviews by Keller4 on crustacean
neuropeptides, Fingerman5. 8 and Waddy and Aiken9
on crustacean endocrine mechanisms, and
Huberman 10 on shrimp endocrinology have been
published. The objective of this review is to
. summarise endocrinological developments in shrimps
which may benefit aquaculture.
Hormonal regulation of growth
Growth pattern in shrimp
All crustaceans have an exoskeleton and ecdysis is
necessary for growth. During early stages of the life
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cycle, shrimps undergo several ecdysis before they
attain juvenile size. As size increases, frequency of
ecdysis is reduced. Female shrimp moult as a
requirement for mating ll . Thus, the control of growth
is dependent on the control of moulting. The moulting
process in crustaceans has been demonstrated I2-ls ; the
process is regulated through ecdysteroid honnones
from the Y-organs during proecdysis and moult-
inhibiting honnone (MIH) produced by a group of
neurosecretory cells (NSC) in the X-or~an sinus gland
complex of the eyestalks of shri mp I3.16- 3.
Role of Y-organs
Gabe24 observed a Y-organ in 58 species of
malacostracan crustaceans. Since then, Y-organs have
been identified and described in numerous species of
crustaceans including shri mps 2s-28. The Y-organ is
generally located in the musculature of the anterior
branchial chamber and appears as a compact mass in
crabs or a less compact mass in crayfish and
lobsters 19. The size of the Y-organ is found to be 60-
300 11m in isopods and amphipods and 1-3 mm in
decapods. Histological studies revealed that the cells
of the Y-organ show secretory activity in relation to
the moult cycle28 .
The endocrine function of the Y-organ in control of
moulting was suggested by Gabe24-29and subsequently
this was demonstrated through surgical experiments
by Echalie~0.31 in Carcinus maenas, and by Passano
and Jyssum32, Maissiae 3-34 and Blanchees in several
other species of decapods, isopods and amphipods
respectively. The first evidence of Y-organs
210 INDIAN] EXP BIOL, MARCH 2005
controlling the moulting process through production
of ecdysteroid hormones was demonstrated by
injecting exogenous ecdysteroid (20-hydro-
xyecdysone) 17. In vitro culture of the Y-organ
provided direct evidence that it produces ecdysteroids
in Procambarus clarkU36 and Penaeus vannamei37
The major secretory product of the Y-organs is an
ecdysteroid which is synthesized from cholesterol 19
and converted to 20-hydroxyecdysone by several
tissues in crustacea. Kaeuser et al. 38 reported lack of
evidence on identification of the target organs of
ecdysteroids and the hormone receptor complex,
which results in increased RNA synthesis through
certain genes. However, recently Durica et al. 39 while
studying the characterization of crab (Uca pugilator)
homologs of ecdysteroid (ECR) and retenoid-X
(RXR) receptors during limb regeneration and oocyte
maturation found that ovarian tissue is a potential
target for hormonal control in crustacea. High levels
of ecdysteroid in blood and tissue during premoult
and reduced levels of same just prior to ecdysis have
been reported by Snyder and Chang40 while working
on the American lobster, Homarus americanus. Blais
et al. 38 reported that in P. vannamei at late premoult
ecdysteroid concentrations increase in the
haemolymph following activation of the Y-organ
while the concentration remains low during postmoult
and intermoult. The rate of ecdysteroid synthesis is
generally controlled by MIH from the X-organ sinus
gland complex. Experiments with eyestalk ablated
shrimp and with Y-organs incubated in vitro in
presence of sinus gland extract indicated that the
X-organ regulates the synthesis of ecdysone in the
Y-organ. It has been reported Palaemon serratus and
Sicyonia ingentis have low levels of ecdysteroids in
eggs at extrusion and the levels increase towards
hatching41 . Alpheus heterochaelis exposed to a
micromolar concentration of 20-0H-ecdysone for 5
days showed that the winter moult cycle was
shortened by 18 days or by 65%42. Vijayan et al. 28
demonstrated that the size and tinctorial affinity of
Y-organ cells in F. indicus change during the moult
cycle. Further, they have shown that the Y-organ
removal in this shrimp inhibits the onset of premoult.
Chan43 cloned a eDNA from the shrimp Metapenaeus
ensis which encodes a nuclear receptor, homologous
to the insect ecdysone-inducible E 75 gene. The
amino acid sequence has all five domains of a nuclear
receptor and it is expressed in the epidermis, eyestalk
and nervous tissue of premoult shrimp.
Role of the eyestalk X-organ sinus gland complex
The eyestalks and their involvement in moult
control were initially discovered by Zelen/.
Hanstrom44 provided a detailed structural description
of the X-organ. With the advancement of science in
neuroendocrine aspects of crustaceans, many workers
have described the X-org~n-sinus gland complex that
contains neurosecretory cells, in details 8,12,17.45,46,
Several types of neurosecretory cells in the X -organs
containing neurosecretory granules in their cell
bodies, axons and terminals have been mentioned
by a number of workers using light and
electronmicroscopl7,48,17.49.46,50. The distribution of
neurosecretory cells in different parts of the optic
ganglia has also been observed and depending on the
location, the terminology adopted was the medulla
terminalis ganglionic X-organ (MTGX), medulla
externa ganglionic X-organ (MEGX) and medulla
interna ganglionic X-organ (MIGX). The sinus gland
is actually formed of axon endings of neurosecretory
cells and glial cells lining haemolymph sinuses 51 .
Exocytosis appears to be the dominant mechanism for
the release of granules into the haemolymph 52 ,17,53,51.
In recent years, Huberman 10 reviewed the chemical
and functional aspects of various ' crustacean
hormones and emphasized their importance in shrimp
aquaculture.
Functional aspects
The role of the optic ganglia of the eyestalk in the
control of various physiological processes including
growth and moulting in crustaceans has been
described54,55,46.4,23. It was demonstrated that eyestalk
ablation in decapods leads to precocious moult, and
moulting occurs several times following ablation,
particularly in young animals. Reimplantation or
injection of eyes talk extract in several crustaceans has
been shown to reverse or inhibit the process of
moulting 13 . The impact of eyes talk removal on
acceleration of the moulting/growth process is due to
loss of MIH, which is predominantly present in the
optic ganglia, particularly in younger stages of the life
cycles. The presence of MIH inhibits the moult
promoting effect of Y-organs 12,52,23.
Once the inhibitory effect of the eyestalk (MIH) on
the moulting process was confirmed in several
decapods, research was conducted to determine the
chemical nature and other physiological functions of
this hormone. MIH was then isolated and its peptidic
nature established in several brachyuran and macruran
 DIW AN: CURRENT PROGRESS IN SHRIMP ENDOCRINOLOGY
species56 . In Carcinus maenas MIH was
characterised57 , purified58 and also sequenced59 . MIH
has also been sequenced in Homarus americanus60
and in Callinectes sapidui l • Aguilar et al. 62 isolated
MIH from the eyestalk of the crayfish Procambarus
bouvieri and compared its sequence with four other
known peptides from H. americanus, C. maenas,
C.sapidus and P. vannamei. The lengths of these
peptides vary between 72 and 78 residues and their
molecular masses between 8 and 9 KDa. The first
cloning of cDNA encoding a MIH like neuropeptide
from P. vannamei was accomplished by Sun63 .
Further, it has been reported that MIH-like mRNA is
found exclusively in the medulla terminalis
ganglionic X-organ MTGXO of the eyestalk and in
the brain the MIH-like gene transcript was detected in
the neurosecretory cells64 • This could mean that a
MIH-like neuropeptide could have specific functions
in the nervous system in addition to its hormonal
function. This neuropeptide has similarities in its
amino acid sequence with other neuropeptides
synthesized in the X-organ, i.e., the crustacean
hyperglycemic hormone (CHH) and the vitellogenesis
inhibiting hormone (VIH). A smaller neuropeptide
(53-55 amino acids) with MIH activity was isolated
from P. bouvien-65. This peptide was later sequenced
and bioassayed62 . Yang et al. 66 isolated and sequenced
a peptide with MIH activity from sinus glands of the
shrimp Marsupenaeus japonicus. In vitro experiments
demonstrated that this peptide inhibited the synthesis
of ecdysone in Y-organs of the crayfish P. clarkii.
Four peptides with both MIH and CHH activity were
isolated from the sinus gland of P. vannamei by
Sefiani et al. 67. It was hypothesized that different
types of peptides that are detected in sinus glands
display different activities, including the site
responsible for MIH activity; but this hypothesis must
be confirmed experimentally 10. Mattson and
Spaziani 68 have shown that MIH is able to inhibit
ecdysteroid secretion by Y-organs in vitro in a dose
dependent manner. The injection of serotonin (5-0H-
tryptamine) lowers the haemolymph ecdysteroid
levels and these effects are revoked by eyes talk
ablation. In this experimental evidence Spaziani
et al. 69 proposed a cycle of MIH regulation sensory
input via 5-HT that would release MIH and inhibit
ecdysteroid synthesis and release by the Y-organs,
and that MIH release would be subjected to a negative
feedback by elevated ecdysteroid titers. Increase in
the mitotic activity in the Y -organ cells after eyestalk
211
ablation in Pachygrapsus mannoratus70 clearly
indicated the possibility of enhanced steroidogenic
activity. Mattson and Spaziani68•71 found that sinus
gland extracts suppress Y-organ steroidogenesis. A
conceptual model defining the mode of action of sinus
gland extracts on steroidogenesis by the Y-organ was
proposed by Mattson and Spaziani72 and the same was
later reviewed by many other workers73 .19 . According
to this model MIH increases intracellular cAMP,
which activates a protein kinase that leads to
inhibition of protein synthesis and steroidogenesis. Gu
et al. 74 while working on MIH in the shrimp
Metapenaeus ensis showed that specific cells in three
different clusters of the X-organ, the sinus gland and
axonal tract of the eyestalk contain MIH. To test the
moult inhibiting activity of MIH, shrimp of intermoult
stage were injected with r-MIH (recombinant protein)
and there was a significant increase in the moult cycle
duration for the shrimp74.
Mandibular organs
The histology and ultrastructure of mandibular
organs (MO) were described in a number of species of
crustaceans25 .9. The main characteristics of cells of
MO are the presence of a centrally located nucleus
with abundant cytoplasm, a large number of
mitochondria and smooth endoplasmic reticulum.
Three types of cells have been identified in MO of
H. americanus75 . The detailed structure of MO of
F. indicus has been described by Vijayan and
Diwan76 .
There is sufficient evidence that terpenoids such as
methyl farnesoate (MF) and farnesoic acid (FA), both
secreted by the MO, have stimulatory effects on
Y-organs and may influence reproductive activity.
Methyl famesoate has been detected in the MOs of
Libinia emarginata77 -79 and several other crustacean
species 8o . The possible implications of MO and MF in
the control of moult and growth have also been
suggested I5-23 . The nature of the MO inhibiting factor
has been explored81 -83 . Eyestalk ablation causes
hypertrophy of the gland and a higher density of
smooth endoplasmic reticulum 84,85. Tsukimura and
Borst86 observed increased levels of MF following
eyes talk ablation in H. americanus. Ablation of MO is
difficult but transplantation of the MO reduces the
length of the moult cycle in Penaeus setiferui7.
While studying regulation of the crustacean MO Borst
et al. 88 described the role of MO in controlling
reproduction and development. Further, they
212 INDIAN J EXP BIOL. MARCH 2005
mentioned 89 that methyl farnesoate (MF) production
by MO is negatively regulated by neuropeptides from
the sinus gland in the eyes talk. , Two neuropeptides
(MO-IH-I and 2) have been isolated from the SG of
the crab, Cancer pagurus that inhibit MF synthesis by
MO of female crabs in vitro. It was revealed that the
regulation of MO is complex and may involve several
SG compounds 89 • Further studies are needed to
determine the exact role of MO in relation to growth
and reproduction.
Hormonal control of reproduction
Reproductive processes of crustaceans are
controlled by hormones. Hormones that influence
gonadal development and spawning are produced in
the brain and thoracic ganglia. The ovaries also
produce hormones. Y-organs, MO and androgenic
glands also produce hormones which promote
gonadal development directly or indirectly. In recent
years, the roles of neuroregulators have been
appraised in enhancing maturation and the spawning
process23 ,90
Reproduction pattern in shrimps
Shrimps are bisexual and development of gonads is
a slow process, which takes place along with the
moulting process during a large part of the adult life.
During reproductive cycles the gonads undergo a
sequence of major morphological and physiological
transformations which require a large amount of
energy. In females, development of the oocyte starts
with the process of oogenesis and in males with
spermatogonial cells. The various stages in the
development of the oocytes as well as spermatozoa
have been descn'b ed lor &
severaI ' 91-93,46.50
speCIes
Determination of sexual maturity of live shrimps in
early stages is difficult. However, as a shrimp
advances towards the ripening phases of gonadal
development, identitification of the maturation phase
is possible. For example, the developing ovary and
developed thalycum of female shrimp for storing
spermatozoa can be seen through the cephalothoracic
region. In males, the development of white
glitterhening spermatheca or sperm boxes are visible
on the bases of the fifth per~iopods. Many have
developed immunochemical techniques for determing
the scale of maturity. Derelle et al. 94 developed an
ELISA titration assay for measuring vitellogenin
synthesis in the freshwater prawn Macrobrachium
rosenberg ii, using a monoclonal antibody. Similar
studies have been done with the lobster
H. americanus95 and crab Callinectes sapidus96 • The
most accurate method now available is based on a
micro-anatomical survey of the reproductive organs
using histological procedures.
Many shrimps are seasonal breeders and in tropical
countries generally the breeding season lasts from
October to February. Environmental factors
particularly temperature, photoperiod and salinity
play an important role in enhancing breeding and
spawning activitl7 . All the marine shrimps breed and
spawn in deep sea waters. The larvae migrate to
shallow coastal waters for their growth to attain the
adult size. Shrimps undergo breeding and spawning
several times and produce millions of larvae during
their lifetime98 • Currently, technologies are available
for domestication of shrimps in hatcheries and
laboratory conditions so that induction of breeding
and spawning at a desired time is possible. The role of
the central nervous system and its coordination with
neuroendocrine centres, along with environmental
cues, are now well understood so far as regulating the
reproductive organs is concerned7 .
Regulation of reproduction
The reproductive activity of crustaceans is partially
under the control of neuroendocrine factors. Panouse 3
showed the presence of gonad inhibiting
hormone(GIH) in the eyes talk of a female prawn,
P. serratus. The increase in ovarian growth after
eyes talk removal was primarily due to removal of the
inhibitory effect on vitellogenesis 46 . Otsu99 showed
the existence of the gonad stimulating hormone
(GSH) in the brain and thoracic ganglion. Based on
these findings it was deduced that gonadal maturation
in shrimp was regulated by two antagonistic
neurohormones, GIH and GSH. This model concept
has been well reviewed I 00,87,97,46,7,23,8. Khalaila et al. 101
while studying the role of eyes talk-borne hormones
on spermatogenic activity in the testis and androgenic
gland of the crayfish, Cherax quadricarinatus, found
that the sinus gland directly controls the activity of the
androgenic gland which suggests an endocrine axis-
like relationship between the sinus gland and AG and
the male reproductive system in decapod crustaceans.
Eyestalk X-organ sinus gland complex
The X-organ-sinus gland complex, which is the
prime neuroendocrine centre in the eyestalks of
shrimps, produces hormonal factors (neuropeptides /
neurohormones) that control the physiological process
of gonads7 • GIH was localized by specific antibodies
 DIW AN: CURRENT PROGRESS IN SHRIMP ENDOCRINOLOGY
in the X-organ SG complex of the lobster
H.americanus,102 prominently at the metanauplius
stage lO3 . By using the technique of nonradioactively
labeled cDNA probes it was possible to detect
neuropeptides in X-organ SG complex of this lobster
and its larvae, which has the property of GIH I04 •105 .
This study revealed that GIH is also present in males.
In the prawn P. varians and the crayfish P. bouvieri
neuropeptides extracted from the eyes talks have a
negative effect on the growth of vitellogenic
oocytes l06.107 Now the eyestalk ablation technique is
being practiced commercially for inducement of
maturatlon. an d spawnlllg
. 0 f sh nmps
' 10846109
" . The
presence of GIH in embryos and larvae \05 may be an
indication of an inhibitory role before adolescence.
Chang et al. I \0 isolated and purified the peptide from
sinus glands of shrimp S. ingentis which is
responsible for inhibition of ovarian development and
spawning. Quackenbush and Keeley III have also
isolated a factor from eyestalks of the shrimp P.
setiferus, which has inhibitory effect on vitellin
synthesis. Similar, studies have been carried out by
Quackenbushl12 in Penaeus vannamei. Huberman lo
recently reviewed the biomolecular aspects regarding
the work carried out on GIH in crustaceans in general
and shrimps in particular. Edomi et at. 113 while
studying GIH of the Norway lobster (Nephros
norvegicus) reported that the GlH is actively involved
in gonad maturation process and plays a more compex
role in control of reproduction and mOUlting. With a
combination of the reverse transcription-polymerase
chain reaction (RT-PCR) and rapid amplification of
cDNA ends approaches, they determined the cON A
sequence of N. no rvegicus prepro-GIH. The
possibility of the involvement of neurohumoral agents
in relation to control of reproduction was debated for
quite some time, but in recent years experimental
evidences are available that 5-HT that is present in the
nervous system including the X-organ sinus gland
complex has a stimulatory effect on reproductive
activity.
Cerebral and thoracic ganglia
A considerable amount of research has been
conducted in crustaceans that indicates the presence
of gonad stimulating factors in cerebral and thoracic
ganglia. Implantation experiments carried out by
Otsu and Gomez and Nayar l14 indicated that the
99
thoracic and supraoesophageal ganglia can induce'
ovarian growth. Injections of aqueous extracts
213
prepared from thoracic ganglia and brain as well as in
vitro incubation experiments indicated vitellogenesis
. .III several crustaceans4511511646117.50
can be stlmulated . . . .
Induction of ovarian maturation in P. vannamei has
been accomplished by injecting lobster brain
extract I 18. By injecting thoracic ganglion extract of
P. semisulcatus, Shoji 50 maturated and spawned
P. monodon. Yano1l9.120 modeled the mode of action
of hormones related to maturation. According to him
GnRH is produced in the brain and in response to this
the thoracic ganglion produces gonadotropins (GnH)
and under the influence of GnH maturation is
effected. However, the molecular structure of GSH is
yet to be elucidated.
Role of endocrine organs
Mandibular organ(MO)
The possible role of the MO in reproductive
activity of crustaceans was reviewed by Laufer
et al. 121 and Waddy and Aiken9. Terpenoids like
methyl farnesoate (MF) and farnesoic acid (FA), both
secreted by MO, influence reproduction of both in
male and female crustaceans. Liu and Laufer found
l22
that the activity of MO is regulated by sinus gland
neuropeptides. It was mentioned that MO-inhibiting
hormonal compounds have the similar type molecular
masses and amino acid composition as noticed with
other sinus gland neuropeptides. Chang et at. 110 by
using a radiolabeled photoaffinity analog of MF found
in S. ingentis the presence of MF binding proteins in
the ovaries, testis and accessory glands in addition to
the haemolymph. The culturing in vitro of ovarian
tissue of the shrimp, P. vannamei in presence of MF
resulted in a significant increase in the size of the
oocytes. This can be interpreted as the involvement of
MF in the early events related to secondary
vitellogenesis. MF was reported to increase fecundity
. cu Iture d sh nmp
III ' P. vannamel·123 .124 . G unaward ene
74
et al. while working on functions and cellular
localization of farnesoic acid O-methyl transferase
(FAMeT) in the shrimp, Metapenaeus ensis reported
that FAMeT directly or indirectly through methyl
farnesoate (MF) modulates the reproduction and
growth of shrimp by interacting with the eyes talk
neuropeptides as a consequence of its presence in the
neurosecretory cells of the X-organ-sinus gland.
Androgenic gland
In decapod crustaceans, the androgenic glands are
generally found associated with the terminal portion
 214 INDIAN 1 EXP BIOL, MARCH 2005
of the male gamete duct. Earlier investigators have
noted the differences and / or uniformities in the
function of the androgenic gland among various group
of crustaceans in general and hermaphroditic and non-
hermaphroditic decapods in particular. Mohamed and
l25
Diwan described the structure of the androgenic
gland in F. indicus and also demonstrated the impact
of bilateral andrectomy on sex reversal of the shrimp.
It was reported that andrectomized male shrimps have
lost their secondary sexual characters and exhibited
absence of sperm in the lumen of their testicular acini.
Decapod androgenic glands are not necessary for
completion of spermatogenesis and their absence
results only in a reduction of spermatogenesis
intensity 125. Mohamed and Diwan l25 reported that the
lack of androgenic glands in F. indicus appeared to
inhibit spermatogonial differentiation. Many studies
have been devoted to determine the chemical nature
. of the androgenic hormone. Laufer and Landau 126
have mentioned that these glands are capable of
producing several compounds including proteins and
the terpenes, hexahydroxy farnesylacetone and
farnesy lacetone, and the exact role of these
compounds is not yet known. Charmantier et at. 22
reported that the androgenic gland hormone regulates
the spermatogenic activity in the testis and is
responsible for the development and maintenance of
the secondary sexual characteristics in male
crustaceans.
V-organs
The role of the Y-organs in crustaceans in the
regulation of the moulting process and growth is well
known 30,127,32,128,33,34,27,23 . However, I'ts ro Ie m .
2o
reproduction has been reported by Chang . As stated
earlier, the Y-organ is the source of ecdysteroids and
correlations between vitellogenesis and ecdysteroids
levels in haemolymph have been reported in some
. 129,87 . The ro Ie 0 f ecd ysteroI'd'
speCIes s m th iattOn
e regu '
of vitellogenesis in female crustaceans is not known.
Ecdysteroids may directly or indirectly participate in
the regulation of spermatogenesis 130 and induction of
gonadal growth in males 121 •
Ovaries
The ovarian tissue in most crustaceans, particularly
in decapods produces vitellogenin-stimulating-
ovarian-hormone (VSOH) and under the influence of
this, the growth of oocytes takes place I31 ,132,10I,92,97.
Besides VSOH, many ecdysteroids have been
identified in the follicle cells and oocytes of the
ovarian tissue21 . Yano l20 described in detail the
control of the process of vitellogenesis by the
endocrine system in penaeid shrimp. He reported that
estradiol-17~ is effective in increasing serum Vg in
the Kuruma prawn, M. japonicus. The hormone
estradiol-17~ is generally distributed in the ovary of
crustaceans and it was suggested that estradiol-17{3
secreted from ovarian follicle cells induces Vg
synthesis in the ovary as a Vg-stimulating ovarian
hormone in penaeid shrimpl20.
Chemistry of hormones
Ecdysteroids
Ecdysteroid or ecdysone, a moulting hormone,
similar to the hormone found in insect prothoracic
glands is produced by the Y -organs in crustaceans 133.
It was also demonstrated that 3-dehydroecdysone is
produced by the Y-organ in several crustaceans
including the shrimp P. vannameP7. Besides,
3-dehydroecdysone, the Y-organ also produces
20-hydroxyecdysone, 25 deoxyecdysone l34 and
several other ecdysteroids 135 • However, the
biosynthetic pathway for the production of ecdysone
is not known, Cholesterol is tr~recusor for ecdysone
p
production, Cholesterol can not be synthesized by
crus taceans 136·
, It must b e 0 b' tame d from dIetary
.
sources. A number of studies indicated the
involvement of ecdysteroids not only in the moulting
process but also in the reproduction of crustaceans 137.
Neuropeptides
Considerable research on chemical aspects of
crustacean neuropeptides has been conducted. Chang
et at. 60 reported an amino acid sequence of lobster
MIH. It has been observed that the amino acid
sequence of lobster MIH was almost identical to that
of crustacean hyperglycemic hormone A (CHHA)
discovered by Tensen et at 138 • Aguilar et at. 62 isolated
MIH from crayfish and compared its sequence with
other four known peptides of crustaceans. Their
lengths vary between 72 and 78 residues and their
molecular masses between 8 and 9 kDa. All have six
cysteines that farm three disulfide bonds. Sun 63 by
using molecular techniques investigated the chemistry
of MIH like neuropeptides of the shrimp P. vannamei.
She reported that this MIH consists of a 72 residue
mature peptide and a 30-residue region of a
propeptide, By using similar techniques Aguilar
et at. 139 isolated the MIH and found that it consists of
a 72-residue peptide with amino and carboxyl termini
 DIWAN: CURRENT PROGRESS IN SHRIMP ENDOCRINOLOGY
and six cysteines forming three disulfide bonds.
Similar investigations have been made by Yang
et al. 66 and it was reported that M. japonicus MIH
consists of a 77-residue peptide with both free amino
and carboxyl termini. Sefiani et al. 67 isolated an MIH-
like peptide from P. vannamei and by mass
spectrometry its molecular mass was estimated to be
8627 Da and consists of only 38 residues. Gu and
Chan43 have isolated MIH from M. ensis and detected
the amino acid sequence of 77 residues preceded by a
signal peptide of 28 residues. Kawakami et al. 140
chemically synthesized a moult inhibiting hormone
(pre-MIH) from the American crayfish, Procambarus
clarkii, which consist of 75 amino acid residues. This
product they found was almost similar to natural pre-
MIH chemically.
Chang 141 reported that activity of growth hormone
peptide has not been reported in either CHH or MIH
assays. Further it is described that the CHHlMIHlVIH
peptide family appears to be a novel group of
crustacean neuropeptides l41 • Much amount of work
has been done on the expression of the CHH gene(s).
Soyez et al. 142 isolated a 7500 Da peptide from the
sinus gland of H. americanus and assayed its GIH
activity in vitro in a shrimp by measurement of oocyte
diameter. They 106 further investigated the structure of
GIH and found that this peptide consists of 77
residues and its molecular weight · is 9135kDa.
Dircksen et al. 143 while studying the crustacean
hyperglycemic hormone (CHH) and CHH-precusor-
related peptides from pericardial organ (PO)
neurosecretory cells in the shore crab, Carcinus
maenas, observed PO-CHH is a 73 amino acid
peptide with a free C-terminus. PO-CHH and sinus
gland CHH have been found to share an identical
N-terminal sequence at positions 1-40 but the
remaining sequence, positions 41-73 or 41-72, differs
considerably. They have also reported that PO-CHH
may have different precursors and CHH genes coding
for precusor products are presumably modified at the
post-transcriptional or post-translational level. It has
been also mentioned that PO-CHH unlike SG-CHH,
has neither a hyperglycemic effect nor it is active in
inhibition of ecdysteroid production by crab
Y-organs.
Katayama et al. 144 studied the structure of moult
inhibiting hormone (MIH) from the Kuruma prawn,
Marsupenaeus japonicus. It has been reported that the
amino acid sequence of MIH is similar to that of
crustacean hyperglycemic hormone (CHH). Further, it
215
was shown that the homology-modeh!d struCture of
M. japonicus CHH was similar to the structure of
MIH with the exception of the absence of the
N-terminal a-helix and the C-terminal tail. This is the
first report on the tertiary structure analysis of a
crustacean neuropeptide. This kind of elucidation is
expected to provide new insights not only on
structure-activity relationship but on the molecular
evolution of the CHH family peptides.
MF is a secretory product of the MO in crustaceans
and is related to insect juvenile hormone a
sesquiterpene that has a significant effect on growth
· ·III·msects. L aUier
an d repro d uctlOn &
et a.I 78 .79·ISO Iated
MF from the haemolymph of crabs. Later a number of
workers have isolated MF from a number of other
crustaceans l41 • Wainwright et al. 82 and Liu et al. 83
have characterised mandibular organ inhibiting
hormones (MOIH) from crabs and found that the
hormones have the molecular masses of
approximately 8400 Da and the amino acid
composition of this neuropeptide is similar to those of
MIH and CHH. There is another compound called
farnesoic acid secreted by cultured MO from P.
clarkU 145 and Cancer magister l46 • Chang 141 has
mentioned that MF is converted rapidly to farnesoic
acid by esterases that are present in tissues.
Androgenic hormone
Once the physiological function of androgenic
hormone was known efforts were made to purify and
characterise the active factors present in this hormone.
Hasegawa et al. 147,148 purified two proteins,
androgenic gland hormone I (AGHI) and AGH II
consisting of 157 and 160 amino acids respectively
with molecular weights of 17.0 and 18.3 kDa. The
other biological active factors isolated were identified
as the terpenes farnesylacetone and hexhydroxy
farnesylacetone l49 • However, the relative role of these
proteins and terpenes is unknown.
Roles of neuropeptides I neurotransmitters in
reproduction
The known neurotransmitters in crustaceans are
acetylcholine (Ach), gamma-aminobutyric acid
(GABA), glutamate, octopamine (OA), dopamine
(DA), and 5-hydroxytryptamine/serotonine (5-HT).
Among the neurotransmitters tested for possible roles
in crustacean reproduction and molting, 5-HT, OA,
and DA seem to be attracting more attention than the
others. Each of these is present in the crustacean
nervous systems.
216 INDIAN J EXP BIOL, MARCH 200S
S-Hydroxytryptamine (S-HT, Serotonin)
5-HT is an ubiquitous substance found in plants
and animals. The Fa1ck-Hillarp fluorescence
histochemical technique for monoamines was used by
several investigators to study the distribution of 5-HT
in crustacean tissues. With this method, 5-HT emits
yellow light. Osborne and Dando '50 detected
fluorescence characteristic of 5-HT in some cell
bodies and in most of the neuropile in the
stomatogastric ganglion of the lobster Homarus
vulgaris.
In recent years, immunocytochemical techniques
have largely supplanted the FaIck-Hillarp
fluorescence technique for localization studies of
biogenic amines '51 . In the eyestalk, immunoreactive
neurons are present in three of the four optic ganglia
(medulla externa, medulla interna, and medulla
terminalis). Some of the immunoreactive cell bodies
in the medulla terminalis send out fibers through the
optic peduncle that terminate in the protocerebrum.
Immunoreactive cell bodies are also present in the
brain. The brain and every ganglion in the ventral
nerve cord displayed at least one immunoreactive cell
body. In addition, several axons in the pericardial
organs are immunoreactive, but the origin and
termination of these axons could not be determined.
In addition to identification and localization of
5-HT by histochemical methods, precise
measurements of tissue concentration of biogenic
amines are now possible. With the HPCL technique,
5-HT was detected in the central nervous system and
hemolymph of P. leniuscu I us. 152 A senes . 0f
experiments done elsewhere revealed that 5-HT
stimulates ovarian development when injected into
the fiddler crab and red swamp crayfish. These crabs
showed increased, dose dependent ovarian
8,1 51
development . indicated that both CHH-related neuropeptides may
Supporting evidence for this hypothesis was
obtained by determining the effects of two 5-HT
agonists on ovarian development in the crab. For this
purpose, the 5-HT releaser fenfluramine and the 5-HT
potentiator fluoxetine were used. The ovaries of crabs
that received fenfluramine, fluoxetine, 5-HT alone,
5-HT plus fenfluramine, or 5-HT plus f1uoxetine
exhibited significant increase in ovarian index and
oocyte size compared to the ovaries of untreated
initial control crabs and saline-injected concurrent
control specimens 151. Supporti ng evidence for a
neurotransmitter role of 5-HT in stimulating GSH
.. . 1153
release in P. clarkii was reported b y SaroJIDI et. a
In M. japonicus, V g synthesis in ovarian pieces
incubated with thoracic ganglion pieces prepared
from vitellogenic females can be stimulated by
5_HTI20.
In addition, 5-HT stimulates testicular maturation
in P. clarkii as shown by the use of a series of 5-HT
agonists and antagonists. 5-HT and its agonists induce
testicular maturation and help in the development of
the androgenic glands. In contrast, 5-HT antagonists
had no stimulatory effects on the testes or androgenic
glands. This stimulatory action of 5-HT on the testes
and androgenic glands was hypothesized to be
indirect, i.e. 5-HT stimulates the release of GSH,
which in tum activates the androgenic glands to
synthesize and release androgenic gland hormone, and
the androgenic gland hormone then triggers testicular
maturation90 . While discussing 5-HT modulation of
crustacean hyperglycerine hormone (CHH) secretion
by isolated cells of the crayfish retina and optic lobe,
Escamilla-Chemal et al. 154 found that retinal CHH
secreting cells correspond to a population of retinal
tapetal cells and optic lobe CHH-secreting cells
correspond to two sUbpopulations of CHH of medulla
terminalis-X organ cells. Further, they mentioned that
CHH secretion generally increases as a function of 5-
74
HT concentration. Gu et al. reported the cloning and
characterization of the cDNA and the gene encoding
the hyperglycemic hormone (CHH-B) of the shrimp
Metapaneus ensis. It was shown that the amino acid
sequence of the hyperglycemic hormone of M. ensis is
identical to that of the CHH-like neuropeptide
(CHH-A) of M. ens is. They further observed the
presence of CHH-B in the eyestalk of vitellogenic
females. At the middle stage of gonadal maturation,a
minimum level of CHH-B transcript and a maximum
level of CHH-A transcript were detected which
play an important role during the female gonad
maturation cycle in shrimp.
Dopamine (DA)
Dopamine (DA) is present in crustacean nervous
systems. Histochemical studies employing the
fluorescence method of Falck and Hillarp indicated
that catecholamines, including DA, are present and
catecholamines ex h1'b'It a green f1 uorescence 151 .
However, both DA and norephinephrine (NE) evoke
this green fluorescence, both being catecholamines.
Much green fluorescence was apparent in the medulla
 DIWAN: CURRENT PROGRESS IN SHRIMP ENDOCRINOLOGY
externa and medulla interna of the eyes talk, and in the
l55
brain and ventral nerve cord of Astacus astacus •
As with 5-HT, immunocytochemical procedures
have been used to demonstrate the presence of DA in
crustaceans. By use of an anti-DA antibody, the
presence of DA-like neurons in the terminal
abdominal ganglion, intestinal nerve, and axons in the
hindgut musculature of Orconectes limos us was
demonstrated. Mercier l56 identified two immuno-
reactive neurons in the abdominal nerve cord of
P. clarkii that contribute axons through the intestinal
nerve to the plexus that surrounds the hindgut. HPLC
analysis confirmed the presence of DA in this nerve 8 .
Role of dopamine in reproduction
DA, when injected into female P. clarkii, inhibited
ovarian maturation. The DA-injected individuals had
a smaller mean ovarian index than the control. The
crayfish given DA plus 5-HT had a significantly
smaller mean ovarian index than the crayfish that
received 5-HT alone, but larger than the crayfish that
received DA alone. 5-HT exerts its effect on the ovary
indirectly, but stimulating GSH release, the inhibitory
action of DA on ovarian maturation induced by 5-HT
could have been due to (1) inhibition of GSH release,
thereby directly counteracting the action of 5-HT; (2)
stimulation of release of the GSH antagonist, GIH; or
(3) both (1) and (2)153.151.
Octopamine (OA)
The presence of OA in central nervous organs of
H. american us was shown by using thin layer
chromatography. With the use of radiolabeled
compounds and HPLC it was shown that OA is
synthesized in the brain of Orconectes. By HPLC, OA
is detected in the eyestalks, brain and hemolymph of
P. leniusculus. Additionally, by HPLC, Butler and
l57
Fingerman found OA in the central nervous system
of U. panacea and Luschen et al. 158 found OA in
Y-organs of C. maenas. Wallace et al. 159 by using
various staining techniques such as treatment with
neutral red and osmium tetroxide and performing OA
analysis by use of a radiolabeled OA precursor
reported the presece of OA in the specific neurons of
the lobster nervous system. Many chromatographic
and electrophoretic techniques were able to identify
neurons in the second thoracic ganglion of
H. americanus that appeared to be the ones that
contain this biogenic amine. The role of OA in
reproduction appears to be at least in part stimulation
217
of contraction of the ovarian walls, liberating the
oocytes.
Red pigment-concentrating hormone (RPCH)
RPCH was isolated from eyestalks of the prawn,
Pandalus borealis. It was later found to be an
octapeptide. Extracts of all portions of the central
nervous system, including the brain and thoracic
ganglia of the crayfishes Cambarellus shufeldi and
Faxonella clypeata have concentrated red pigment
hormone in the integumentary chromatophores,
thereby revealing that RPCH is widely distributed
throughout the central nervous system of the
crayfish l61 .
Role of RPCH in reproduction
RPCH, in addition to its hormonal role in
regulating pigmentation, appears to have another
hormonal role , stimulation of MF synthesis in the
mandibular organ of P. clarkii. MF may function as a
hormone involved in the regulation of crustacean
reproduction to complement the two peptidergic
neurohormones, GIH and GSH, that have well
documented roles in controlling gonadal maturation in
crustaceans 136.
Opioid peptides
The first report of the presence of an opioid-like
substance in crustaceans was published by Mancillas
et al. 161. Through immunocytochemistry, they found
Leu-Enk-like immunoreactivity in all the retinular
cells of the spiny lobster, Panulirus interruptus, and
P. clarkii. In addition, such immunoreactivity was
also apparent in nerve fibers in chiasma 3 that run
from the medulla interna to the medulla terminalis.
Later, other crustaceans were similarly studied.
Immunoreactivity was apparent in the retinular cells,
lamina 'ganglionaris, sinus gland, optic peduncle, the
three chiasma, and medulla terminalis. Immunoreac-
tivity was present in ovarian follicle cells and
avitellogenic and early vitellogenic oocytes but not in
fully ripe oocytes. In the testes, the immunoreactivity
was present in spermatogonia, spermatocytes and
spermatids but not in mature spermatozoa. This was
the first demonstration of opioid-like Immunoreac-
tivity in a crustacean gonad.
Role of opioids in reproduction
The potential involvement of an endogenous opioid
system in the regulation of ovarian development in
218 INDIAN J EXP BIOL, MARCH 2005
U. pugilator has been investigated in viV0 162 • Injection
of synthetic Met-Enk into female crabs significantly
slowed ovarian maturation. The inhibition was dose-
dependent. The mean ovarian index and mean oocyte
diameter of the crabs that received the opioid were
significantly smaller than the corresponding values
for the saline-injected concurrent control specimens.
In contrast, injection of the opioid antagonist,
naloxone, produced dose-dependent ovarian matura-
tion. It was hypothsized that the opioid (1) stimulates
GIH release, (2) inhibits GSH release or (3) does both
(1) and (2).
The role of an opioid in testicular maturation of the
fiddler crab was also investigated. Injection of
synthetic Met-Enk into male crabs significantly
slowed maturation of the testes. The inhibition was
dose-dependent. The mean testicular index, testicular
weight, testicular lobe diameter and the number of
mature spermatocytes in each testicular lobe were
significantly less than the corresponding values for
the saline-injected concurrent control crabs. Injection
of naloxone produced dose-related testicular
maturation 162.
Conclusion
The shrimp industry today all over the world is
developing rapidly, particularly in Southeast Asian
countries due to the high demand of shrimp in the
international market and also due to its high export
value. The success of the industry often depends on
the availability of the shrimp resource on a continuous
basis. Today the shrimp industry is mostly dependent
for this resource from the wild and to meet the
demand, shrimps are being exploited continuously
and in the process overexploitation takes place, and as
a result shrimp catches have dwindled and in some
places even the fishery has declined to a large extent.
In order to reverse this trend and manage this resource
on a sustainable basis the FAO has recently
introduced the Code of Conduct For Responsible
Fisheries (CCRF) for strict adoption by the all the
maritime countries. Even if this code becomes
effective, it will take a long time to rejuvenate the
depleted stock. Therefore, in order to fulfill the
demand of shrimps by the industries on a continuous
basis, one of the alternate ways is to produce them in
captive conditions.
In the recent past several efforts have been made to
evolve methods for the induction of maturation and
spawning of shrimps in captive conditions. Though
eyes talk ablation has been accepted as the most
successful method for induction of maturation and
spawning, this method does not work for many
shrimps of commercial value. Moreover, there are
other inherent and social problems in adopting this
technique. Hence, the need of the hour is to explore /
devise methods involving other possible neuro-
endocrine manipulations. The use of neurohumours or
neurotransmitters, or extracts from the brain and
thoracic ganglia containing reproductive hormones,
has to be explored to initiate gonadal maturation.
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