Aquaculture Research, 2006, 37, 1429^1436

doi:10.1111/j.1365-2109.2006.01578.x

Panagrellus redivivus mass produced on solid media

as live food for Litopenaeus vannamei larvae
Ulfert Focken1, Christian Schlechtriem1,, Matthias von Wuthenau1, Armando Garc| a-Ortega2,
Ana Puello-Cruz2 & Klaus Becker1
1

Aquaculture Systems and Animal Nutrition in the Tropics and Subtropics (480B), University of Hohenheim, Stuttgart,

Germany
2

CIAD (Research Centre for Food and Development), Aquaculture and Environmental Management, Mazatlan, Mexico

Correspondence: U Focken, Aquaculture Systems and Animal Nutrition in the Tropics and Subtropics (480B), University of Hohenheim
(480B), 70593 Stuttgart, Germany. E-mail: focken@uni-hohenheim.de
Present address: Institute of Aquaculture, University of Stirling, Stirling, Scotland, UK.

Abstract

Introduction

The free-living soil nematode Panagrellus redivivus is

well known to be an excellent food source for rst
feeding sh larvae. It represents an alternative to the
highly expensive Artemia, which is commonly used.
The lack of a proper method for mass production of
P. redivivus has prevented its wider use in commercial
hatcheries. A new cultivation method allows the production of a sucient quantity of nematodes to deliver a standardized and permanently available live
food of high quality, throughout the larval rearing
period. In two experiments ^ carried out at the Centro de Investigacion en Alimentacion y Desarrollo,
Mexico ^ several feeding regimes were established to
prove the quality of the mass produced P. redivivus for
larvae of Litopenaeus vannamei, the Pacic white
shrimp. Two dierent nematode treatments were
compared with a no-feed group and a control group
that was fed with Artemia. All treatments had an additional algal co-feed and were run in ve replicates.
Panagrellus redivivus was cultured on two dierent
media (wheat/corn our and oat our) to compare
these for their suitability as high-quality live food for
the larvae. Shrimp fed nematodes grown on wheat/
corn medium reached the postlarval stage earlier
than those from other treatments. The nematode
treatments showed promising results; however,
further research is needed on the development of improved culture media or enrichment methods to
further increase the nutritional value of P. redivivus.

The contribution of the crustacean to the world

aquaculture production values (excluding algae)
reached 22.6% in 2004. Shrimps ^ representing 83%
of crustacean production in tonnes and 85% in values ^ are the largest group of species in crustacean
farming. Most of these shrimp species derive from
the family Penaeidae. Litopenaeus vannamei contributes to 47% of aquacultural shrimp production and
43% of production values (FAO 2006).
One of the major bottlenecks in aquaculture production is the rearing of sh and crustacean larvae
(Lavens & Sorgeloos 1996). This is due to the fact that
many of the species cultured depend on live
food during larval stages (Sorgeloos, Dhert & Candreva 2001). This live food should be easily available, reproducible and economical (Watanabe & Kiron1994).
Problems occurring in supply with this live food may
prevent successful larval rearing, which limits the
whole production system (Guillaume, Kaushik, Bergot & Metailler 2001).
The most frequently used live food organism is the
brine shrimp Artemia sp. This small crustacean has
the advantage that its culture can be started from
dried eggs (Sorgeloos & Persoone 1975; Liao 1992).
These dormant cysts can be stored for longer periods
in cans and, if needed, used as a convenient o-theshelf live food (Lavens & Sorgeloos 2000). All that is
needed for incubation is the hydration of the cyst in
warm, aerated seawater and illumination. This is sufcient to start the embryonic development in the dormant cysts and leads to the hatching of the nauplii
(Sorgeloos & Persoone 1975).

Keywords: Litopenaeus vannamei, larviculture,

live food, nematodes
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Journal Compilation r 2006 Blackwell Publishing Ltd

1429

Nematodes as live food for shrimp larvae U Focken et al.

Although Artemia is particularly convenient to use

in hatcheries (Wickins & Lee 2002), it also has some
prominent negative aspects. The most common ones
are: high costs, a highly variable hatching rate, quick
growth, the varying nutritional quality and the property to consume algal feed and therefore to compete
with the cultured species for food (Biedenbach,
Smith, Thomsen & Lawrence 1989; Lavens & Sorgeloos 1996, 2000). The main drawback for future use
is that the cyst production in the Great Salt Lake
(Utah, USA) which is the major production site of Artemia, is limited and does not satisfy the growing
world demand (Lavens & Sorgeloos 2000).
Owing to the obvious limitations of Artemia, other
live organisms have been examined for their use in
penaeid shrimp larviculture; copepods, rotifers,
Daphnia, Moina and nematodes have been suggested
by various authors (Wilkenfeld, Lawrence & Kuban
1984; Lavens & Sorgeloos 1996; Guillaume et al.
2001; Lee, OBryen & Marcus 2005).
Samocha and Lewinsohn (1977) reported on the
successful use of the free-living soil nematode Panagrellus redivivus in rearing postlarvae of Penaeus semisculcatus and Metapenaeus stebbingi. The authors gave
nematodes in a ground mixture with Sepia, Artemia,
Tubifex and Enchytraeus. Kahan, Bar-El, Brandstein,
Rigbi and Oland (1980) suggested nematodes as a potential candidate for a live food organism in rearing
sh fry. Their suitable size, high nutritional values
and an easy cultivation promised nematodes to be a
valuable feed. Wilkenfeld et al. (1984) stated that the
nematodes were able to substitute Artemia in penaeid
larval rearing diets. Their experiments showed the
capability of Farfantepenaeus aztecus, Litopenaeus setiferus and L. vannamei to consume and survive on
P. redivivus as the only food source from the Protozoea 1 (PZ-1) stage. Experiments of Biedenbach et al.
(1989) with P. redivivus in the larval rearing of L. vannamei conrmed these results.
Hitherto, nematodes are most commonly cultured
on a variety of solid and liquid media. In these production systems, only small amounts of nematodes
could be produced. The lack of a proper mass production technology for nematodes is the most limiting
factor to commercial application (Fisher & Fletcher
1995) and further investigation on such techniques
is recommended (Biedenbach et al. 1989).
Fisher and Fletcher (1995) produced nematodes
by the means of a biological fermenter lled with
culture medium, inoculated with Escherichia coli
and P. redivivus. Although designed for the production of large quantities of nematodes, it seems that

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Aquaculture Research, 2006, 37, 1429^1436

this system has not yet been commercially

established.
Bedding (1981) and Bedding, Staneld and Crompton (1991) developed a solid-medium technique for
the production of entomopathogenic nematodes like
Neoplectana spp. and Heterorhabditis spp. With the
novelty of the exploitation of the third dimension,
this system enlarges the usable surface inside the
growing system by using crumbled polyether polyurethane sponges to create an interstitial space. This
space allowed optimal reproduction conditions and
served as a living habitat for the nematodes. It also
guaranteed sucient aeration. From the solid-medium system of Bedding (1981) and Bedding et al.
(1991), Ricci, Fi, Ragni, Schlechtriem and Focken
(2003) developed a system for the mass production
of P. redivivus. It consisted of autoclavable plastic bags
lled with sponges soaked with medium. The bags
were inoculated with Saccharomyces cerevisiae to
guarantee a monoxenic culture. The system was
aerated and kept humid during the 11^13 days of
incubation. Several media, medium quantities and
inoculation densities have been investigated. Compared with the biological fermenter proposed by Fisher and Fletcher (1995), the bag system by Ricci et al.
(2003) delivers higher multiplication factors and is
less complicated to apply (Schlechtriem, Ricci, Focken & Becker 2004a).
Rouse, Webster and Radwin (1992) have shown
that the nutritional character of P. redivivus depends
on the nutritional qualities of the culture medium it
is grown on.Wilkenfeld et al. (1984) used soft dough of
corn our and deionized water; Biedenbach et al.
(1989) worked with a mixture (50/50) of nonbleached wheat our, corn meal and deionized water
to produce soft dough. Kumlu, Fletcher and Fisher
(1998) used baed 250 mL asks and a medium of
animal protein, corn oil and yeast inoculated with
E. coli. Ricci et al. (2003) used an oatmeal-based medium and a soluble medium made up of puried ingredients, resembling the proximate composition of oat.
Additional investigations concerning the eect of an
added oil source (sh oil or sunower oil) on body
composition, average yields and multiplication factors of the nematodes were conducted by Schlechtriem, Ricci, Focken and Becker (2004a, b). The
enrichment of culture media with further lipid
sources (capelin oil, cod liver oil and marilla oil) was
tested by Kumlu et al. (1998).
There are several studies on the use of mass-produced P. redivivus in the rearing of rst feeding sh
larvae (Santiago, Ricci & Reyes-Lampa 2004;

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Aquaculture Research, 2006, 37, 1429^1436

Nematodes as live food for shrimp larvae U Focken et al.

water, resulting in a constant humidity of air. The nematode culture was incubated for12 days in a climatically controlled room at an average of 22 1C.
For harvest, culture media with nematodes were
placed in a sieve clad out with a milk lter
(+200 mm, Hygia Favorit by Paul Hartmann AG,
Heidenheim, Germany). This sieve was placed on a
plate half lled with water. Nematodes crawling
through the milk lter dropped into the water. This
water was ltered several times with a 112 mm plankton net until most of the nematodes were removed.
The mass of nematodes was washed with distilled
water to clean them from adhering culture media or
yeast. Nematodes were harvested daily and stored
until feeding in a Petri dish in a fridge at ca. 4 1C.

Schlechtriem et al. 2004a, b; Schlechtriem, Focken &

Becker 2005). The authors concluded that the bag
system is suitable for the mass production of nematodes as live food for rst feeding larvae of sh and
crustacean species, and Ricci et al. (2003) stated that
it should be applicable for small-scale live feed producers in developing countries due to its simple technology. Although there are several studies of the new
mass production system and it is known that P. redivivus is a suitable food for rst feeding shrimp larvae,
no trials have been conducted to evaluate the use of
mass-produced P. redivivus in the rearing of L. vannamei larvae.
In this study, P. redivivus was mass produced
according to the method of Ricci et al. (2003) on two
types of cereal-based media that are commonly used
for the cultivation of nematodes. The suitability of
both kinds of nematodes was compared with Artemia
in their use as live food for rearing L. vannamei larvae.

Feeding trials
The experimental set-up for shrimp larval rearing
was adapted from Puello-Cruz, Sangha, Jones and Le
Vay (2002). Twenty bottles with a round body and a
long neck were placed in a water bath at 28 1C in an
air-conditioned chamber at 22 1C with a 12 h dark/
light cycle. Each bottle contained 1.5 L of fresh seawater at 35  1ppt. The water was aerated by the
means of plastic tubes (+0.5 cm) ending in a glass
tube, reaching down to the bottom. The current of
the air stream was regulated, so that one air bubble
per second was emitted, which guaranteed sucient
aeration and food distribution in the water column.
Bottles were stocked with batches of 150 nauplii of
L. vannamei. In the rst trial, stocking was performed
using a volumetric method. Even though nauplii at
this stage still deplete internal reserves and do not
feed on algae, a mixture of 20 algal cells mL 1 (70%
Chaetoceros muelleri, 30% Isochrisis galbana) was
added to guarantee feed supply once the animals
reached the protozoea stage.

Materials and methods

Nematode production
The nematodes were mass produced on a monoxenic
solid culture (single microorganism: Saccharomyces
cerevisiae) according to Ricci et al. (2003). Two dierent
culture media were used. One consisted of a mix of
wheat and corn our (50/50) (Biedenbach et al. 1989)
and the other exclusively of oatmeal (Ricci et al. 2003).
The ours were mixed with a 0.8% marine salt solution (Tetra Marin), and crumbled polyether polyurethane sponges (75 g) were added. The mixture of
sponges and culture media was inserted into autoclavable bags (50  30 cm) and then autoclaved for
55 min at 121 1C. Each bag was inoculated with approximately 5.5  105 organisms of P. redivivus. The
bags were aerated via plastic tubes; the incoming air
passed an air lter (0.2 mm) and a bottle lled with

Table 1 Treatments and feeds for each successive stage of larval development of Litopenaeus vannamei
Artemia

Stage

Wheat/corn-nematodes

Oat-nematodes

Nauplius 5

Mixed algae: 20 cells mL

Mixed algae: 20 cells mL

Mixed algae: 20 cells mL

Mixed algae: 20 cells mL

Protozoea 1 Mixed algae: 50 cells mL

Mixed algae: 50 cells mL

Mixed algae: 50 cells mL

Mixed algae: 50 cells mL

Mixed algae: 50 cells mL

Mixed algae: 50 cells mL

Protozoea 2 Mixed algae: 50 cells mL 1

P. redivivus: 75 mL 1 (100 mL
Protozoea 3 P. redivivus: 75 mL
to Mysis 3

(150 mL

Mixed algae: 50 cells mL 1

) P. redivivus: 75 mL 1 (100 mL

) P. redivivus: 75 mL

(150 mL

Algae-only

) Artemia: 210 mL

Mixed algae 50 cells mL

Number of nematodes fed in the second experiment are given in parenthesis.

P. redivivus, Panagrellus redivivus.

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Nematodes as live food for shrimp larvae U Focken et al.

1432

(ANOVA) and the Tukeys test after testing for deviations from normal distribution by the using Kolmogorov^Smirnovs test. The dierences were reported
as statistically signicant when Po0.05.

Results
In the two experiments conducted, the most striking
dierence was observed in the rates of development
and survival. In the rst experiment, all animals of
the Artemia treatment were found in the postlarval
stage at day 11. An average of 16% of the shrimp fed
oat-nematodes transformed into postlarval stages.
No further postlarvae were found in any other treatment. (Fig. 1).
In the rst experiment, the highest average survival was found in the oat-nematode treatment (85.8%),
followed by shrimps from algae-only treatment
(72.6%), the Artemia treatment (70.0%) and the
wheat/corn-nematode treatment (69.6%). No signicant dierence in survival could be observed between the treatments from the rst trial (Table 2).
The average dry weight of the shrimps from the Artemia treatment (90.9 mg) was signicantly higher than
that of shrimps from the other treatments (Table 2).
The weight of shrimps fed oat nematodes (45.5 mg),
wheat/corn nematodes (37.9 mg) or algae-only
(30.3 mg) did not dier signicantly from each other.
Shrimps fed Artemia (5.7 mm) were signicantly
longer than the animals from other treatments.
Shrimp from the wheat/corn-nematode treatment
Postlarvae

Mysis 3

Mysis 2

100

75
%

Four dierent treatments with ve replicates each

were tested (Table 1): one Artemia treatment, two nematode treatments (oat and wheat/corn) and one
treatment of algae only. In the rst stages ^ from Nauplii 5 to Protozoea 3 (PZ3) ^ all treatments were fed
with an algae mixture of 70% C. muelleri and 30%
I. galbana. From stage PZ2 on, nematodes were given.
Owing to its size, Artemia could only be fed from stage
PZ3 on. This kind of feeding had been used as Artemia treatment in former experiments in CIAD. To the
remaining ve replicates, no further food but algae
was added.
The mixture of algae was prepared from algae
(C. muelleri and I. galbana) strains regularly used in
CIAD. Algae were cultivated in aerated seawater
(35  1ppt) at a temperature of 22  1 1C. The numbers of cells were evaluated under the microscope,
the mixture was prepared and the appropriate
amount of algal solution (Table 1) was injected into
the bottles by means of a syringe.
Artemia cysts (Premium Brine Shrimp eggs, Prime
Artemia, Midvale, UT, USA) were incubated for 24 h at
28 1C, an average salinity of 35 ppt and constant illumination. The hatched Artemia nauplii were separated
from the remaining cysts, whether unhatched or broken, and inserted into the bottles after counting.
Nematodes were constantly extracted and counted
daily. The number of nematodes per millilitre in the
resulting solution was calculated and the appropriate
amount of animals was injected into the bottles. The
number of nematodes and Artemia and the amount of
algae fed were adapted towards the growing demands of the growing larvae (see Table 1). In the rst
experiment, 75 nematodes mL 1 were given for each
developmental stage. In the second experiment,
100 nematodes mL 1 were given for PZ3 and
150 nematodes mL 1 for the subsequent stages. In
this experiment, the nauplii of L. vannamei were
counted individually. Apart from these changes, the
set-up of both experiments was identical. Representative samples of nematodes and Artemia were freeze
dried for later chemical analysis.
The experiments were terminated when 90% of
the shrimps in one replicate of one treatment reached
the rst postlarval stage. The larvae and postlarvae
were counted for survival, development stages were
evaluated and the length of the animals was measured. The accumulated average dry weight of the
larvae was measured after freeze-drying.
Statistical analysis was performred with the programme GRAPHPAD PRISM 4.03 and INSTAT 3.05. Data
were analysed using a one-way analysis of variance

Aquaculture Research, 2006, 37, 1429^1436

50
25

0
Artemia

Oat Nem

WC Nem

Algae

Diet

Figure 1 Average number of shrimps found in the dierent developmental stages in the dierent treatments from
the rst experiment (Day 11). Oat Nem, nematodes reared
on oat medium; WC Nem, nematodes reared on wheat/
corn medium.

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Aquaculture Research, 2006, 37, 1429^1436

Nematodes as live food for shrimp larvae U Focken et al.

Table 2 Survival rate, nal dry weight and length of Litopenaeus vannamei from Experiment 1
Diet

Survival (%)
Dry weight (mg)
Length (mm)

Artemia

Oat nematodes

Wheat/corn nematodes

Algae

70.0  34.6
90.9a  15.6
5.7a  0.1

85.8  9.8
45.5b  18.3
4.6bc  0.2

69.6  18.6
37.9b  8.2
4.8b  0.2

72.6  17.9
30.3b  4.0
4.4c  0.1

Mean  SD, ve replicates per treatment.

Means not sharing a common superscript are statistically dierent at Po0.05.

(4.8 mm) were signicantly longer than shrimp algae

only (4.4 mm); shrimps fed oat nematodes (4.6 mm)
were intermediate and did not dier from wheat/corn
nematode or algae treatment.
In the second experiment, in which the feeding
intensity for the nematode treatments had been
increased, the development in the wheat/corn nematode treatment was faster. On day 9, in some replicates of this treatment, all animals had reached
postlarval stage, the average was 74%, followed by
the oat-nematode treatment with 70%. Only 4% of
the shrimp from the Artemia treatment had transformed to the postlarval stage at that day (Fig. 2).
Survival rates, dry weight and length at the end of
the second experiment are given in Table 3. Shrimps
fed algae-only, Artemia and wheat/corn-nematodes
had a survival rate of 90.4%,85.1% and 74.8% respectively.The lowest survival rate was observed in the
oat-nematode treatment (37.1%). The survival of
shrimps fed oat-nematodes was signicantly lower
Postlarvae

Mysis 2

Mysis 3

Mysis 1

100

75

50

25

0
Artemia

Oat Nem

WC Nem

Algae

Diet

Figure 2 Average number of shrimps found in the dierent developmental stages in the dierent treatments from
the second experiment (Day 9). Oat Nem, Nematodes
reared on oat medium; WC Nem, Nematodes reared on
wheat/corn medium.

than the survival rate of shrimps fed Artemia and algae-only. The survival rate of animals fed wheat/
corn-nematodes was not statistically dierent from
those of any other treatment. In dry weight, no signicant dierences occurred between shrimps fed
with Artemia (82.1 mg) or nematodes, whether these
were grown on oat (82.8 mg) or wheat/corn (80.2 mg)
medium. The weights of the shrimps from these three
treatments was signicantly higher than that of
shrimp from the algae-only treatment (26.8 mg). In
the second experiment, the average length of
shrimps fed oat nematodes or wheat/corn nematodes
was identical (5.2 mm). The dierence from shrimps
fed Artemia (4.8 mm) was not signicant. All three
treatments showed a signicant dierence in length
compared with the shrimps from the algae-only
treatment (3.7 mm).

Discussion
The experiments ended when 90% of the shrimps of
one replicate of a treatment had transformed into the
rst postlarval stage. Shrimps in the other treatments
found in larval stages ^ for example mysis ^ were
judged to show a slower development compared with
those already in the postlarval stage. During the two
experiments, it was observed that if shrimps of one
treatment performed well in terms of fastness of development, the survival rate of the same treatment
was lowered. This may be due to the higher number
of moults that postlarvae have already performed
during their development compared with the lower
stages. Generally, the mortality in this experiment
was similar to that observed by Puello-Cruz et al.
(2002) in the same set-up (47.6^70.0%), but somewhat lower than the values observed by Wilkenfeld
et al. (1984) and Biedenbach et al. (1989), who report
survival rates between 73% and 90%. The large standard deviation indicates that survival may have also
been aected by factors not related to the feeding

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Nematodes as live food for shrimp larvae U Focken et al.

Aquaculture Research, 2006, 37, 1429^1436

Table 3 Survival rate, nal dry weight and length of Litopenaeus vannamei from Experiment 2
Diet

Survival %
Dry weight (mg)
Length (mm)

Artemia

Oat nematodes

Wheat/corn nematodes

Algae

85.1a  8.6
82.1a  8.8
4.8a  0.3

37.1b  33.8
82.8a  17.4
5.2a  0.3

70.8ab  30.3
80.2a  11.9
5.2a  0.3

90.4a  3.0
26.8b  1.2
3.7b  0.2

Mean  SD, ve replicates per treatment.

Means not sharing a common superscript are statistically dierent at Po0.05.

treatments, oering scope to improve survival in all

treatments. To reach postlarval stages faster means
to have shorter intermoult periods in that the animal
can replace energy losses during moults. Moulting
animals are also especially vulnerable to cannibalism (Wickins & Lee 2002) and weak in the end of
moulting (Hartnoll 1982). The whole progress of
moulting and emerging from the old exosceleton is
an energy-consuming activity (Hartnoll 1982) and
all of this may weaken and stress the shrimp larvae
and could lead to a lower survival rate.
Hartnoll (1982) described the growth of crustacean species as a discontinuous process with successive moults and intermoults. During the moults, the
animals shed their old integument and gain body
length and mass until the new integument has hardened (Wickins & Lee 2002). No major growth in
fresh weight occurs in the intermoult, which is the
time between two moults. Owing to the hard exoskeleton, the growth is restricted and may be ignored for
most purposes (Hartnoll 1982). The discontinuous
growth pattern may also explain the lack of a signicant dierence in the average dry weight as observed
between shrimps from the dierent treatments of the
second trial. Shrimps fed oat-nematodes were signicantly longer than shrimp from the Artemia treatment and therefore a higher dry weight should be
expected. However, the growth occurring during
moulting is mainly due to the absorption of water; later, this water will be replaced gradually by proteins
(Wickins & Lee 2002). Just having passed into the
postlarval stage, this water has not been replaced by
proteins and dry weight of the freshly moulted
shrimps may be lower than that of those still in the
last larval stage.
The gradual increase in nematodes injected into
the bottles during the second experiment induced a
similar performance of larvae fed nematodes compared with shrimps fed Artemia. Shrimps from both
nematode treatments outperformed those from the
algae-only treatment in all relevant criteria except

1434

survival rate. The signicantly (Po0.05) lower survival rate in the oat-nematode treatment during the
second experiment may be due to the faster development of the shrimp larvae or to lower water quality, a
topic that is most critical during larvae culture
(Wickins & Lee 2002). In the experiment conducted
no water exchange was performed to guarantee an
undisturbed larval development. However, for feeding high numbers of nematodes, a water exchange
seems crucial, due to nematodes dying and decaying
after 72 h in salt water (Biedenbach et al. 1989). The
introduction of residues of nematode culture medium into the experimental system may also have a
detrimental eect on the water quality. The problem
of a lower water quality using nematodes as live feed
was described by other authors (Wilkenfeld et al.
1984; Biedenbach et al. 1989; Santiago et al. 2004).
However, washing nematodes even more thoroughly
before being fed to shrimp larvae may reduce the risk
of introducing micro-organisms or residues of culture medium. A regular water exchange or adding
water during larval culture ^ as practiced in commercial hatcheries ^ should further enhance water
quality. In future experiments, higher survival rates
may be accomplished if these points are taken into
consideration.
Litopenaeus vannamei has a dietary protein requirement of about 20^30% (New 1980); however,
their larvae, that are more carnivorous, may have a
higher requirement (Lee, Smith & Lawrence 1984).
The body compositions of nematodes produced on
oatmeal and wheat/corn medium were described in
detail by Schlechtriem et al. (2004a) and Biedenbach
et al. (1989). Nematodes grown on oat and wheat/corn
medium had a crude protein content of 62% and 48%
in the dry matter respectively. The amino acid composition of both types of nematodes was similar to
that of plankton and Artemia and can thus be considered to be appropriate to cover the essential amino
acid requirement of shrimp larvae. The lipid requirements of penaeid larvae are not fully understood but

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Aquaculture Research, 2006, 37, 1429^1436

a crucial role of polyunsaturated fatty acids, cholesterol and phospholipids became apparent during the
last years (Jones,Yule & Holland1997; Guillaume et al.
2001). As other animals, crustaceans are incapable of
producing linoleic acid (LIN, 18:2n6) and a-linolenic
acid (ALA, 18:3n3), but they can use these as precursors to synthesize the corresponding higher polyunsaturates. However, the ability for bioconversion of
ALA and LIN to highly unsaturated fatty acids
(HUFA) as eicosapentaenoic (EPA; 20:5n3) and decosahexaenoic (DHA; 20:6n3) is limited (Teshima, Kanazawa & Koshio 1992) and therefore requirements
for these compounds must be satised by the diet.
Phospholipids, which play an important role in cellular membranes as well as sterols that cannot be
synthesized by crustaceans, are further essential
dietary compounds. The chemical composition of nematodes can be signicantly inuenced by the composition of the culture medium, that is at least partly
ingested by the nematodes. The lipid class composition of P. redivivus mass produced on dierent media
was described in detail by Schlechtriem,Tocher, Dick
and Becker (2004c). Nematodes produced on cerealbased medium are poor in EPA and contain no DHA;
however, Schlechtriem et al. (2004b) described that
the addition of sh oil to culture media can increase
the HUFA content of the nematodes signicantly.
The results of this study show that mass-produced
P. redivivus seems to be a potential replacement forArtemia in the larval rearing of L. vannamei. However,
further studies are required to test whether tailoring
the body composition of P. redivivus towards the specic need of crustacean species can further improve
the nutritional value and thus the suitability of nematodes in larval rearing of shrimps.

Acknowledgment
This work was partly funded by a grant of the Eiselen
Foundation Ulm to MvW.

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