Broodstock Nutrition And
Management In Crustacean
Kumar Meena*., Pronob Das1., Md. Shahbaz
Akhtar2., Sekar M1.,and
*Central Inland Fisheries Research Institute, Barrackpore,
Kolkata, 700 120
1. Central Institute of Fisheries Education, Mumbai, 400 061
2. Directorate of Coldwater Fisheries Research, Bhimtal, 263
author: Dharmendra Kumar
* Email : firstname.lastname@example.org
future of aquaculture production, as in other animal production
systems, is towards greater control of the physical, chemical and
biological variables surrounding the production system. The trend
from collection of wild fry to the development of hatcheries for fry
production, from the use of wild spawners to the use of maturation
systems and ultimately closing the reproductive cycle to use
domesticated and selected stocks is clear. At the same time, there is
an increasing trend towards production efficiency and
cost-effectiveness, not least in the hatchery stage, particularly
where the supply — demand situation progresses from quantity
production to quality production. These trends are becoming more
evident in all forms of commercial aquaculture. Broodstock and larval
nutrition are key elements underpinning this progress towards greater
control and domestication. However, there is substantial work to be
done if rearing of aquaculture species is to approach the level of
control and understanding which are evident in the poultry, swine and
ruminant sectors. Research into shrimp broodstock nutrition is
gaining importance with the increasing use of domesticated and
genetically selected stocks for aquaculture. Also,given the high cost
of feed in a captive broodstock maturation facility, estimated at
around 50% of total cost, as well as a heavy reliance on fresh feed
items such as squid, mollusk meals and polychaete worms which may
vary in quality and availability, there is a demand for feeds
specifically formulated for use in maturation diets. It has also been
pointed out (Doyle, personal communication) that the development of
genetically improved lines of shrimp may benefit from a simultaneous
development of feeds specifically tailored to individual strain
During penaeid shrimp
maturation, nutrient reserves, mainly from the hepatopancreas, are
mobilised to support ovarian and testicular maturation, gametogenesis
and vitellogenesis (Harrison, 1990; 1997). Tissue reserves in the
hepatopancreas can be depleted rapidly so that the diet becomes the
most important contributor of nutrients to the developing egg.
This is particularly true
when, as is the case in most captive maturation facilities, eyestalk
ablation is used to accelerate the process of gonadal maturation. The
hormonal and metabolic changes that come around during such forced
maturation may take place when the nutrient reserves are insufficient
to support rapid ovarian development placing an even larger burden on
the diet as a source of essential nutrients.The nutrition of penaeid
shrimp broodstock has recently been reviewed (Wouters et al., 2001a).
The authors recognize that there is a dearth of information in the
scientific literature on the subject, possibly due to the expense and
complexity of running sufficiently rigorous nutrition experiments on
shrimp broodstock. In addition, much of the research has been carried
out with fresh feeds, either alone or in combination with formulated
diets or dietary supplements.
to the importance of lipids in crustacean maturation, much of the
work carried out to date has focused on this aspect, particularly the
requirement for highly unsaturated fatty acids (HUFA) and
phospholipid. During maturation, lipids are mobilized from the
hepatopancreas in many species and dietary lipids rapidly processed
for transport to the developing ovaries.Total lipid does not appear
to be important although Wouters et al. (2001b) reported that
excessively high total lipid in the diet had an adverse effect on
ovarian maturation and feed consumption, possibly due to satiation.
However, the average lipid level in commercial broodstock diets (10%)
appears to be around 3% higher than in grower feeds used in
commercial culture ponds.
Highly unsaturated fatty
acids (HUFA), especially 20:5n-3 and 22:6n-3, are abundant in ovarian
tissues and are believed to be an important component of live and
formulated maturation diets. Diets deficient in n-3 HUFA have been
found to have a negative effect on ovarian development, fecundity and
egg quality (Wouters et al., 1999a).Arachidonic acid (20:4n-6; AA)
has been detected at high levels in the ovaries of wild shrimp and is
also abundant in some of the best fresh feed items such as
polychaetes (bloodworms), clams and mussels (Harrison, 1997). The n-6
HUFA are known to be precursors of the prostaglandin hormones, which
act in reproduction and vitellogenesis. According to Wouters et al.
(2001a), formulated maturation diets appear to be deficient in AA as
well as relatively low eicosapentanoic acid (EPA) levels. It has been
proposed that the ratio of n-3 to n-6 levels in the diet is important
and that it should be around 2—3 to 1 (Wouters et al.,
1999b).Phospholipids, mainly phosphatidylcholine and
phosphatidylethanolamine appear to be predominant in the shrimp ovary
and there seems to be a requirement for phospholipids in the diet.
Several studies (Ravid et al., 1999; Wouters et al., 1999b) have
demonstrated the effects of phospholipid levels in the diet and it
has been suggested that broodstock diets should contain more than 2%
phospholipid to ensure that 50% of total egg lipid is in this form
(Cahu et al., 1994).
Cholesterol is the
precursor of steroid hormones and it is known that shrimp have a
requirement for cholesterol in the diet. Cholesterol is stored in the
hepatopancreas and is mobilized during maturation. Some of the live
feed organisms used in maturation diets have relatively high
cholesterol levels (e.g. squid, clams) although to date there has
been limited research into the effects of dietary cholesterol on
maturation and reproduction (Wouters, 2001).During maturation, the
level of triacylglycerides (TAG) in the ovaries increases as they are
incorporated into the egg and decrease after spawning (Ravid et al.,
1999; Wouters et al., 1999b). Triacylglycerides appear to be the
principal energy source in eggs and nauplii and their importance in
reproduction, and egg and postlarval quality has been shown (Palacios
et al., 1998; 1999).
AND AMINO ACIDS
Maturation is a time of
intense protein synthesis and it is likely that the requirement for
protein is higher at this time (Harrison, 1997). Wouters et al.
(2001a) report that the protein content of formulated diets in their
studies was around 50% but that this was still low compared to the
level in fresh feeds.
Some studies have shown
changes in protein content of the ovaries associated with egg
development and spawning, and with spawning success. Harrison (1997),
found an increase in ovarian protein levels associated with ovarian
development followed by a sharp decrease after spawning in the shrimp
Paratelphysa hydrodromaus and several authors have noted a similar
increase in farmed penaeid shrimp (Castille and Lawrence, 1989). A
marked difference has also been noted in the protein content of the
hepatopancreas and ovaries of wild and domesticated females of
Litopenaeus vannamei with good repeat spawning performance which have
been found to have significantly higher protein content than females
with poorer spawning performance (Palacios et al., 2000).
Carbohydrates do not
appear to be essential for shrimp broodstock diets although Palacios
et al. (1998; 1999) related egg glucose levels with larval quality
and broodstock condition. Carbohydrates can be used as cost-effective
ingredients to contribute to glycogen accumulation in the
hepatopancreas as well as providing other benefits in the broodstock
diet, acting as binders and possibly playing a role in transport of
nutrients in the hemolymph (Harrison, 1997).
vitamin and mineral requirements for shrimp broodstock diets are
relatively unknown with only a few studies on vitamins A, C and E.
Alava et al. (1993) found that ovarian maturation was slower when fed
a diet deficient in either vitamins E, A and C. Vitamin E appears to
be important in crustacean broodstock nutrition.Chamberlain (1988),
reported a correlation between vitamin E deficiency and the
percentage of abnormal sperm in Litopenaeus setiferus and Cahu et al.
(1991) found an improvement in hatching rate with increasing dietary
vitamin E correlated to increasing—tocopherol levels in the
egg.Wouters et al. (1999b) found a similar correlation to that
observed by Cahu et al. (1991) between spawn and hatch quality with
a-tocopherol levels in wild spawners and nauplii of L. vannamei.
They found that mature
ovaries and nauplii contained higher levels of —tocopherol than
immature and spent ovaries. Harrison (1997) also speculated that
vitamin E in the egg yolk may also act as a natural antioxidant.
(1997), suggested the importance of dietary vitamin A due to its
accumulation in the ovaries of crustaceans during maturation. Vitamin
C content of eggs of Fenneropenaeus indicus are also influenced by
the levels in the diet, and high hatching rate has been related to
high ascorbic acid levels in the eggs (Cahu et al., 1995). Harrison
(1997) assumes that vitamin D is important in broodstock diets due to
its probable role in calcium and phosphorus metabolism in
crustaceans.Harrison (1990) discussed the possibility that mineral
deficiencies or imbalances could have a negative impact on crustacean
reproduction and may play a role in oocyte resorption, reduction in
reproductive performance and egg quality. Studies into mineral
requirements are rare due to several complications (Wouters et al.,
2001a). Where studies have been conducted, diets were formulated with
mineral mixes with added calcium, phosphorus, magnesium, sodium,
iron, manganese and selenium (Marsden et al., 1997; Xu et al., 1994).
Spent broodstock of L.
vannamei had lower levels of calcium and magnesium in the muscle and
lower magnesium levels in the hepatopancreas (Mendez et al., 1997),
possibly due to a combination of dietary deficiencies and losses
through moulting and transfer to the eggs. Copper also decreased in
the hepatopancreas, possibly through transfer to the ovaries,
although it increased in the muscle tissue. It is clear that more
studies need to be undertaken into mineral nutrition in broodstock
synthesise carotenoids de novo, and a dietary source of these
pigments is required. During sexual maturation, most crustaceans
accumulate carotenoids in the hepatopancreas and during
vitellogenesis, these are transported in the hemolymph as
carotenoglycolipoproteins to accumulate in the eggs as part of the
lipovitellin protein. Dall (1995) found that free astaxanthin levels
in the developing ovaries of Penaeus esculentus increased from 2 to
34 ppm and in the digestive gland, from 20 to 120 ppm.
astaxanthin, are strong antioxidants and probably play a role in
protecting the broodstock nutrient reserves and developing embryos
from oxidation (Dall et al., 1995; Merchie et al., 1998). It has also
been suggested (Harrison, 1997) that they act as pigment reserves in
the embryos and larvae for the development of chromatophores and
eyespots, and as a vitamin A precursor (Dall, 1995).
Hunter (1999) found that astaxanthin supplementation (50 mg/kg) of
the diet resulted in increased egg production in Penaeusmonodon but
could not demonstrate any benefit of astaxanthin supplementation on
either hatching rate or metamorphosis to zoea 1 stage.
It has been suggested
that some of the more successful live feed organisms may provide
benefits through the provision of hormones or their precursors.
Naessens et al. (1997) speculated that part of the reason for the
success of reproductive adult Artemia biomass supplementation of the
diet of L. vannamei broodstock could be due to the presence of
specific hormones or analogous peptides in the Artemia that provoked
a response in the shrimp. Bloodworms used in maturation have also
been found to contain methyl farnesoate, an ecdysone hormone that has
been shown to increase reproductive performance in the spider crab
Libinia emarginata (Laufer et al., 1987), L. vannamei (Laufer et al.,
1997), P. monodon (Hall et al., 1999) and the crayfish Procambarus
clarkii (Laufer et al., 1998). In P. clarkii, the hemolymph titer
increased from basal levels during early vitellogenesis, peaked
during mid-cycle and then returned to basal levels when the ovaries
were in late vitellogenesis.
Nucleotides, the basic
building blocks of nucleic acids, are recognised as important
elements in mammalian nutrition especially during periods of rapid
growth or physiological stress (Van Buren, 1994) and also appear to
play a key role in the immune system. Traditionally, nucleotides have
not been considered essential nutrients although de novo synthesis
and salvage pathways are thought to be costly processes in metabolic
terms. Several studies have demonstrated that dietary sources of
nucleotides can have beneficial effects and the term 'conditionally
essential' has been used to describe their role in nutrition
(Carver and Walker, 1995).
Exogenous sources of
nucleotides are thought to optimise the functions of rapidly dividing
tissues, such as those of the developing embryo and young, and the
reproductive and immune systems.Most aquaculture diet ingredients of
animal and plant origin contain nucleotides although there are
differences in the concentration and availability. Nucleotide content
is particularly high in ingredients such as fish solubles, animal
protein solubles, fishmeal, legumes (adenine is particularly high in
blackeyed peas), yeast extracts and unicellular organisms such as
yeasts and bacteria that are rich in RNA or DNA (Carver and Walker,
1995; Devresse, 2000).
Devresse (2000) noted
that the low digestibility of whole yeast compared to yeast extract
may be related to the protein (nitrogen) solubility as yeast extract
has much higher protein solubility than whole yeast. He also noted
that, although fish solubles are highly digestible, they leach easily
in water affecting availability.
Reproduction and egg
development have a high requirement for RNA and DNA and it may be
expected that increasing the availability of nucleotides in
broodstock diets may have a beneficial effect on egg development.
Recently, research has demonstrated the effect of a
nucleotide-enriched diet for broodstock nutrition in aquaculture
(Gonzalez-Vecino, 2002; Gonzalez-Vecino et al., 2003).
Nucleotide enrichment of
broodstock diets for Atlantic halibut (Hippoglossus hippoglossus) and
haddock (Melanogrammus aeglefinus) resulted in a general trend
towards better spawning performance and egg quality with the
nucleotide diet. Total egg yield was 30% higher in the halibut fed
with the nucleotide diet and the relative fecundity, mean egg
density, hatching rate and survival of yolk-sac larvae were also
significantly improved. Haddock fed on the nucleotide-enriched diet
also had significantly higher fertilization and hatching rates. To
date, no work has been published on nucleotide supplementation of
broodstock diets for shrimp but it would be interesting to conduct
some trials to determine if broodstock diets enriched with
nucleotides might offer similar benefits in shrimp maturation and
breeding. Similarly, the potential for nucleotide supplementation of
diets for shrimp larvae should also be investigated.
are a fewer information regarding the knowledge of bloodstock and
larval nutrition of crustaceans available which creates many gaps
between researcher and literature . This is partly the result of the
complexity and expense of conducting research into these two areas.
However, given the increasing importance of domestication in shrimp
aquaculture particularly, there is a need for an increased focus on
these areas. The role of nutrition in broodstock and maturation
performance and in increasing larval survival and quality will be
fundamental to obtaining optimal performance from domesticated
stocks. Even in species where domestication is not an issue, the
improvement of maturation performance and larval production remains a
key goal in improving the efficiency of production systems. It is not
clear how far the goal of complete replacement of live feeds may be.
It is likely that this will be attained in broodstock diets long
before larval feeds. Indeed, given the complexities inherent in
supplying a complete nutritional package in a small particle, it may
be that this goal is never reached.
it may be possible to supply a range of diet particles (e.g. high
lipid particles, high protein particles, carbohydrate particles etc.)
that are aimed specifically at providing the right mix of nutritional
elements in the culture tank that will expose the larvae to the
appropriate nutritional mix. Alternatively, it may be that, as
expressed by D'Abramo (2002), the complexity of the ontogeny of
larval nutritional physiology may mean that technical success will be
based on a compromise between the desire to provide a complete diet
package and the need to strive for simplicity in formulation and
manufacture. In this paper maximum emphasis is given on different
sources as nutrition and their beneficial effects on broodstock
quality but very limited information are available regarding this
module. This information will provide a base line for researcher for
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