Aquaculture 315 (2011) 155–166

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Aquaculture
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / a q u a - o n l i n e

Review

Status and recommendations on marine copepod cultivation for use as live feed
Guillaume Drillet a,f,⁎, Stéphane Frouël b, Mie H. Sichlau c, Per M. Jepsen d,f, Jonas K. Højgaard d,
Almagir K. Joarder e, Benni W. Hansen f
a
DHI Water and Environment Pte Ltd, Pandan Loop 200, 128388, Singapore, Singapore
b
DANISCO Animal Nutrition, DANISCO France SAS, Paris, France
c
DTU-Aqua, Technical University of Denmark, Kavalergården 6, DK-2920 Charlottenlund, Denmark
d
AKVA group Denmark A/S, Nyrnberggade 31A, 1., DK-2300 København S., Denmark
e
Department of Communication, Business and Information Technologies, Roskilde University, DK-4000 Roskilde, Denmark
f
Department of Environmental, Social and Spatial Change, Roskilde University, DK-4000 Roskilde, Denmark

a r t i c l e i n f o a b s t r a c t

Article history: Copepods are important crustaceans studied because of their key role in ecology, trophic biology, fisheries
Received 12 August 2009 management, in modeling the flow of energy and matter, ecotoxicology, aquaculture and aquarium trade. This
Received in revised form 17 February 2011 paper discusses various aspects of the state of knowledge of copepod culture at large scales and provides the
Accepted 17 February 2011
scientific community with ideas and concepts that could improve and quicken the development of copepod
Available online 25 February 2011
mass cultures. As a framework for discussion, we use a conceptual scheme from Teece (1988) and adapted it
to our goal: ‘how to capture value from a copepod product’. The suggestions include: 1) optimize cultures by
Keywords:
Copepod culture
automation and implement recirculation technology for improving water quality; 2) use harpacticoid and
Live prey cyclopoid copepods in industries that can produce large amounts of these prey on site at any given time; but
Cannibalism use calanoid copepods for industries limited in production time and those that export copepod products (e.g.
Density eggs); 3) select preferentially local copepod species and if possible species with lipid conversion capabilities;
Population selection 4) optimize sex ratio and selection/cross-breeding to develop suitable copepod strains for aquaculture;
Probiotic 5) explore the use of probiotics for improving the fitness of copepod cultures; and 6) encourage copepod
producers/retailers to use/develop an efficient sales and marketing strategy.
© 2011 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
2. Part 1: Competitive production — biological challenges and perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
2.1. Which copepod to raise?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
2.2. Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
2.3. Density . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
2.4. Egg and nauplii cannibalism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
2.5. Optimization of sex-ratio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
2.6. Selection and cross-breeding to obtain optimized populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
2.7. Enrichment and use of probiotics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
3. Part 2: Complementary technologies — technical challenges and perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
3.1. Copepod cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
3.2. Temperature–salinity–oxygen–light. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
3.3. Water quality, recirculation and automation of cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
3.4. Conservation and shipping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
4. Part 3: Sales and marketing — commercialization strategy for copepod products . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
5. Conclusions and recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

⁎ Corresponding author at: DHI Water and Environment Pte Ltd, Pandan Loop 200, 128388, Singapore, Singapore. Tel.: +65 93266187.
E-mail address: [email protected] (G. Drillet).

0044-8486/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2011.02.027
156 G. Drillet et al. / Aquaculture 315 (2011) 155–166

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

1. Introduction ecotox/ (US Environmental Protection Agency). Furthermore, aqua-

culture and the aquarium trade are in desperate need of copepods to
Free living copepods have been intensively studied because of supply the rapid development of their respective industries. Støttrup
their impact as key players in the marine pelagic environment. In (2000) and Payne et al. (2001) put forward the hypothesis that the
terms of biomass, copepods can represent up to 80% of the inclusion of copepods in aquaculture will enhance the number of
mesozooplankton (Mauchline et al., 1998). They are an important successfully reared fish species. The same hypothesis can be applied to
food source for planktivorous fish and fish larvae in general (e.g. Fox increase the production of various species of ornamental fish for the
et al., 1999; Möllmann et al., 2004). Their life cycles, vital- and aquarium trade which would lead to a decrease in fishing pressure on
physiological rates are intensively studied to understand the effects of natural stocks. The benefits include a reliable delivery of fish for the
various environmental conditions on the pelagic ecosystem. Experi- end-users and a unique chance for protecting delicate environments
mental zooplankton studies have benefited from the small-scale like coral reefs. One of the keys to successful larval finfish rearing is the
cultivation of copepods. Also, when compared to field sampled high nutritional quality of copepods compared to other traditionally
specimens, the advantage is that the history and condition of used live prey (Drillet et al., 2006b). Extensive cultures of copepods have
cultivated copepods are known and can even be manipulated. already been achieved in order to supply aquaculture industries and
Information derived from the experiences and practices of copepod aquarium trade needs with high quality live feed (Engell-Sørensen et al.,
culturing is important for a variety of research areas such as copepod 2004; Evjemo et al., 2003; Olivotto et al., 2008; Rajkumar and Vasagam,
genetics, feeding behavior, population dynamics, parameterization of 2006; Sørensen et al., 2007; Toledo et al., 1999; Van der Meeren et al.,
standing stocks, production rates for ecosystem models, etc. 2008; Wilcox et al., 2006) and systems including the use of copepods
Since the 1960s culturing copepods have become increasingly have been patented (patents numbers: CN1288967 and JP2007044042,
more reliable and approximately 60 copepod species have been as examples).
successfully raised (Mauchline et al., 1998). To our knowledge, the The present contribution evaluates different limitations that slow
oldest copepod culture is from the Danish Technical University of down the development and realization of new cultivation methods,
Denmark (Acartia tonsa) which was isolated from Øresund (a strait while at the same time, proposes possible novel and necessary actions
between Denmark and Sweden) in 1981 (Støttrup et al., 1986). The based on a multi-disciplinary approach. The group of authors
World Copepod Culture Database was initiated in 2006 at Roskilde represents university researchers with backgrounds in zooplankton,
University (http://copepod.ruc.dk/main.htm) in an attempt to supply aquaculture, business and management, and professionals from the
and share knowledge between copepod scientists, aquaculturists, and private sector with expertise in microbiology and aquaculture
the public at large. The database contains details on various cultures consulting.
and up-to-date recent knowledge on cultivation procedures. To date, We present our evaluations and suggestions in three major parts
approximately 30 copepod cultures have been referenced in the dealing with: 1: Competitive production — biological challenges and
database (Table 1). perspectives; discusses the challenges of copepod culture at the
When copepod cultures are required in fundamental research biological level, 2: Complementary technologies — technical challenges
fields such as phylogeny, physiology and ecology, there is a need for and perspectives; provides technical information and innovations about
reliable continuous cultures. This includes the potential availability of copepod cultures, and 3: Sales and marketing — commercialization
copepods for toxicity testing in bio-assays as was demonstrated by status and perspectives; proposes a suite of recommendations which
Karlsson et al. (2006) and Palacios-Caceres and Pereira (1997) with need to be addressed to develop profitable intensive copepod
more references of eco-toxicity studies found at http://cfpub.epa.gov/ cultivation.

Table 1
Extract of the copepod culture list from the World Copepod Culture database (data are from January 2009). (http://copepod.ruc.dk/main.htm Drillet and Dur, 2006–2011).

Genus/species Geographical origin Cultivation conditions, temperature, salinity, light regime, food

Acartia grani Barcelona Harbor (Spain) 19 °C/38 ppt/12L:12D/Rhodomonas salina

Acartia sinjiensis Townsville Chanel, (Australia) 27–30 °C/30–35 ppt/18L:6D/Tetraselmis chuii and T-iso
Acartia southwelli Pingtung (Taiwan) 25–30 °C/15–20 ppt/12L:12D/Isochrysis galbana
Acartia tonsa Øresund (Denmark) 17 °C/30 ppt/dim light/Rhodomonas salina
Amphiascoides atopus USA 25 °C, 12L:12D, cultured phytoplankton
Apocyclops royi Pingtung (Taiwan) 25–30 °C/15–20 ppt/12L:12D/Isochrysis galbana
Centropages typicus Gulf of Napoli (Italy W Mediterranean Sea) 19–21 °C/38 ppt/12L:12D/Prorocentrum minimum/Isochrysis galbana/Tetraselmis suecica
Eurytemora affinis River Seine Estuary (France) 10–15 °C/15 ppt/12L:12D/Rhodomonas marina
Eurytemora affinis Gironde Estuary (France) 10–15 °C/15 ppt/12L:12D/Rhodomonas marina
Eurytemora affinis Loire Estuary (France) 10–15 °C/15 ppt/12L:12D/Rhodomonas marina
Eurytemora affinis Baie de l'Isle Verte/St Laurent Estuary (Canada) 10–15 °C/15 ppt/12L:12D/Rhodomonas marina
Euterpina acutifrons Mediteranea 19 °C/38 ppt/12L:12D/Rhodomonas salina
Gladioferens imparipes Swan River (Perth, Australia) 23–27 °C/18 ppt/continuous dark/T-Iso and Chaetoceros muelleri
Mesocyclops longisetus Florida (USA) http://edis.ifas.ufl.edu/IN490
Microcyclops albidus Florida (USA) http://edis.ifas.ufl.edu/IN490
Oithona davisae Barcelona Harbor (Spain NW Mediterranean) 20 °C/30 ppt/natural light/Oxhyrris
Pseudodiaptomus annandalei Pingtung (Taiwan) 25–30 °C/15–20 ppt/12L:12D/Isochrysis galbana
Temora longicornis North Sea 15 °C/30 ppt/continuous dark/Thalassioiera weissflogii, Rhodomonas salina,
Heterocapsa, Prorocentrum minimum
Temora longicornis Plymouth (Devon, UK) Temp according to current sea temperatures, 30–36 ppt/12L:12D/mixture
of Isochrysis galbana, Rhodomonas and Oxyrrhis
Temora stylifera Gulf of Napoli (Italy W Mediterranean Sea) 19–21 °C/38 ppt/12L:12D/Prorocentrum minimum/Isochrysis galbana/Rhodomonas baltica
Tisbe battagliai Brixham (Devon, UK) ?
 G. Drillet et al. / Aquaculture 315 (2011) 155–166 157

2. Part 1: Competitive production — biological challenges cultures (Phelps et al., 2005; Su et al., 2005). Cyclopoids definitely
and perspectives offer a great potential for aquaculture and aquarium trades but they
have not been intensively studied maybe because of the difficulties in
2.1. Which copepod to raise? harvesting nauplii from culture (Støttrup, 2006) and in the lack of
storage possibilities for eggs. Calanoid copepods are pelagic and
Some copepods have more potential for live feed cultures because present in the entire water column. They have been particularly
of particular life history traits. For instance, a fast reproductive cycle studied in aquaculture because they are natural prey for fish larvae.
and low mortality are two factors that matter in terms of viability of Nauplii of calanoid copepods have successfully been fed to a wide
commercial copepod cultures, but one could also argue for biochem- range of fish species including cod, halibut, flounder and barramundi.
ical composition or/and swimming behavior. The relevant candidates Several calanoid copepod species have been proposed as candidates
for commercial culturing belong to three orders of copepods for which for aquaculture production; and the commonly studied species are
advantages/disadvantages have been reviewed (Støttrup, 2006). In from the following genera Acartia, Eurytemora, Gladioferens, Parvoca-
short, harpacticoid copepods are epi-benthic copepods with in many lanus and Centropages (O'Bryen and Lee, 2005). Advantages from
cases pelagic nauplii and have been proven to have superior some of the calanoids include the capacity of storing eggs (Drillet
nutritional qualities compared to Artemia and rotifers (Cutts, 2003; et al., 2007; Marcus and Murray, 2001) but limitation resides in the
Drillet et al., 2006b). Species of flatfish, gobies, salmonids, ciaenids, difficulty of attaining high density cultures without negative side
and blennies are sometimes considered to be obligatory harpacticoid effects, like a decrease in hatching success or high mortalities (Jepsen
feeders, at least for a period in their life cycle (Coull, 1990). Most et al., 2007; Peck and Holste, 2006). Maximum densities of the
harpacticoids can be grown at high densities (e.g. from 10,000 to cultivated calanoid species proposed for aquaculture purposes are
400,000 ind L− 1, Støttrup, 2003) in high area/volume systems within the range of 100–1000 adults L− 1 (Støttrup and McEvoy,
provided by incorporating 3D structures in the rearing tanks (Fig. 1) 2003), but recent studies have shown small-scale cultures attaining
(Stottrup and Norsker, 1997). However, most harpacticoids are egg densities of 4000 A. tonsa ind L− 1 while keeping a low and density-
carriers (Williams and Jones, 1994) and harvest of nauplii can become independent mortality (Drillet et al., 2006a).
difficult in large systems. This has been recently solved in a semi-
automatic rearing system producing 1–2 million nauplii daily in 2.2. Food
average in a 600 L system (Steenfeldt, 2008) and production systems
that are able to generate up to 8.2 million of Nitokra lacustris nauplii The amount and quality of food are essential parameters that
per day in 266 L systems exist (Rhodes, 2003). Cyclopoid copepods enhance the production of copepod cultures. Effects of food quantity and
with obligatory pelagic nauplii are pelagic and are used occasionally in quality have been assessed in natural populations because of their
aquaculture and densities of ~5000 ind L− 1 are possible to achieve in importance at the ecological level. Most of the information present in

Fig. 1. Schematic ideal large-scale copepod production. Copepod culture for free spawning and egg carrying species are incorporated. Benthic copepods are produced in systems
containing 3D structures. Harvest of nauplii/eggs for further utilization or storage decreases the risk of cannibalism. Recirculation procedures provide high quality water to which
automatic feeding is added. A surveillance system equipped with multi-sensors keeps track of variations in vital parameters in cultures, allowing a minimum of labor force during
production (figure made using Google SketchUp 7™ and MS office™).
158 G. Drillet et al. / Aquaculture 315 (2011) 155–166

the literature can be extrapolated and used for culture purposes. can bio-convert fatty acids, this would permit one to feed them diets
Quantity-wise, all copepods do not have the same need to reach their based on cheap and easily accessible low quality food items like flour
maximum growth and production potentials even when closely or yeast as it was shown by Kahan and Azoury (1981), Kahan et al.
related (Jonasdottir, 1989). Too little food tends to slow down stage (1982) and Takano (1971a,b) for Nitocra spinipes, Gladioferens
development, increase the competition for food between the imparites and Tigriopus japonicus, respectively. These cheap food
individuals, and ultimately increases mortality (Berggreen et al., sources would be of great interest for lowering the cost of copepod
1988). Most of the studies reported focus on the effects of optimal productions; although one of the problems generated by the use of
food concentration at the individual level. However, the improve- flour or yeast as food for copepods in the same manner as used for
ment of copepod cultures increases the competition for food rotifers and Artemia, is that the resulting water quality is often poor.
resources and investigations on optimal food concentration in high Two reports (Chapman, 1981; Gyllenberg and Lundqvist, 1978)
density cultures should be performed (also because phytoplankton proposed that copepods Neocalanus plumchrus and Cyclops oitho-
production is costly). noides could use dissolved glucose as a food source. However, for
Food quality should be adjusted for the target number of copepods, water quality reasons, this would not be an optimal solution for
and the size of the prey should be consistently adapted not only to the feeding copepods in cultures due to expected problems with
species but also to the development stages that need to be fed uncontrolled growth of heterotrophic bacteria and the potential risk
(Berggreen et al., 1988; Hansen et al., 1994). Previous research has for developing pathogenic microorganisms in the system.
shown that a mixture of different diets generally enhanced the
somatic growth and egg production of copepods in the laboratory 2.3. Density
(Harris, 1977; Jonasdottir, 1994; Klein Breteler, 1980), but also the
right food item at the right development stage increases the overall Density of individuals is crucially important to copepod cultures.
success (Koski et al., 2006; Murray and Marcus, 2002). The food items Some copepods are much more sensitive to high density (hundreds
should be adapted to the copepod feeding habits for example whether per liter; Ban and Minoda, 1994) than other live organisms like
the copepods raised are raptorial feeders or suspension feeders. In the rotifers (millions ind L− 1; Moretti et al., 1999). Thus, they have been
case of a raptorial feeder, phytoplankton might not be the only food used only occasionally in fish larval feeding. The density question is of
item that copepod cultures could require. In cultures of the central importance because production costs are directly related to
carnivorous copepod Rhincalanus nasutus, Artemia nauplii mixed possible densities of copepod cultures. Calanoids are considered poor
with diatoms were used with success (Mullin and Brooks, 1967). In candidates for mass cultivation simply because they do not survive
the case of feeding on a non-algal diet (Takano, 1971a, b; Guérin and well at high densities, but harpacticoids and cyclopoids can commonly
Gaudy, 1977), it is relevant that the cultivated copepods also use the be raised at densities reaching a few thousand ind L− 1 or more (see
microbial- and ciliates prey associated with the degradation of the Lee et al., 2005; Støttrup and McEvoy, 2003; Steenfeldt, 2008). The
added algal diet as a food source. The capacity to survive on purely trade off is most often that increased density is negatively affecting
bacterial food was described for Tisbe holothuriae by Rieper (1978), survival, developmental time, growth, reproduction and egg hatching
and Zillioux (1969) described microphageous ciliates as a source of success (Zhang and Uhlig, 1991; Kahan et al., 1988; Ban and Minoda,
food in his Acartia culture system. Ciliates are used as the sole food 1994; VanderLugt and Lenz, 2008); suggesting that a balance must be
source for raising other copepod species (Suárez et al., 1992). The achieved. However, it is interesting that for various species of the
studies by Klein Breteler (1980) give good ideas of the potentials same genus and even populations of the same species, density-
given by heterotrophic dinoflagelates in copepod cultures showing dependent egg production and hatching success showed highly
they may be relevant food items. Nassogne (1970) tested different divergent results (Camus and Zeng, 2009; Jepsen et al., 2007; Medina
food organisms on the development and culture of Euterpina and Barata, 2004; Peck and Holste, 2006). However, these differences
acutifrons, as did Støttrup and Jensen (1990) for A. tonsa and Abu may be due to inherent differences between populations or to
Rezq et al. (1997) for Tisbe furcata. The list of food items relevant for growing conditions as proposed by Drillet et al. (2008b) and therefore
various target copepod species is comprehensive and could be a fast conclusions should be avoided. Nevertheless, high culture
review by itself. In all copepod cultures nuisance organisms like densities increase the intensity of resting egg production of
microphageous and histophageous ciliates, nematodes, and even Eurytemora affinis and Acartia sinjiensis (diapause and delayed
rotifers regularly appear. The potential negative effect these organ- hatching eggs) (Ban and Minoda, 1994; Camus and Zeng, 2009). To
isms exhibit either directly by preying on copepod eggs or indirectly our knowledge, this has never been shown for any other copepod
as food competitors needs to be addressed. Recently a method to species but is well known in other crustaceans (e.g. Daphnia spp.;
eradicate rotifers was proposed by Steenfeldt and Nielsen (2010), and Zadereev, 2007). This density-dependent diapause egg production
abrupt salinity drop is by personal experience effective against seriously raises the question of the viability of intensifying batch
ciliates, but nematode fauna is still difficult to eradicate. However, cultures and possibly that chemical communication between cope-
some microbial components are beneficial as food items for the pods could be a limit to enhancement. On the other hand, this effect
copepods. New possibilities are to establish an equilibrium between could also be investigated in order to deliberately aim at producing
the cultured copepods, their algal food and relevant protozoans. If these particular types of eggs for use in aquaculture. This strategy
successful it will provide the copepods with food supplements to its might allow long-term storage of resting eggs, which opens up new
algal food with a different biochemical composition and prey size. This perspectives for storage and shipping (see Section 2.4). However, if
strategy is most likely more realistic than constantly fighting nuisance accumulation of chemical cues/metabolites in the water can lead to
organisms, but it requires research to achieve this delicate equilibrium detrimental effects on the culture, then changing the water regularly
in the cultures. or even applying water treatment technologies like circulation (see
Biochemically, some copepod species seemingly have the capacity later) allows batch or continuous cultures to reach high densities as in
to bio-convert shorter chains of fatty acids (e.g. 18:3) to long chain the case of calanoids (e.g. N4000 ind L− 1 with 5% mortality day− 1;
poly-unsaturated fatty acids (PUFAs). This has been proposed for Drillet et al., 2006a).
Eucyclops serrulatus and Paracalanus parvus by Desvilettes et al.
(1997) and Moreno et al. (1979). Nevertheless, it has also been 2.4. Egg and nauplii cannibalism
reported that some species of copepods cannot grow well on
phytoplankton lacking highly unsaturated fatty acids (Støttrup and A problem facing dense copepod cultures is cannibalism due to the
Jensen, 1990; Klein Breteler et al., 1999). If some species of copepods increased encounter rates of individuals in the rearing tanks.
 G. Drillet et al. / Aquaculture 315 (2011) 155–166 159

Cannibalism has been reported for many copepod species (e.g. unknown, but there are some indications that they can have a
Sinocalanus tenellus, Centropages abdominalis, Tigriopus fulvus, and reproductive outcome (Miller et al., 2005; Sillett and Stemberger,
Acartia sinjiensis). Often, smaller individuals (younger stages) are 1998; Dharani and Altaff, 2002). Species with intersex may have an
more likely to be the victims to the more developed stages (Daan advantage if they are able to reproduce successfully, because they are
et al., 1988; Ohman and Hirche, 2001; Ohno and Okamura, 1988; Uye able to secure the population in a highly patchy planktonic
and Liang, 1998). Hada (1991) and Hada and Uye (1991) showed for environment and thereby give a higher reproductive outcome. As
S. tenellus that the ingestion rate of their own nauplii increased with long as we do not know if the intersex species have a high successful
increasing nauplii density to an asymptotic value and that cannibal- reproductive outcome, it is of interest to keep good food quantity to
ism took place even when alternative food was available. When adult prevent sex changes from occurring. However, if natural conditions
females occurred at a density of 10 ind L− 1, the mortality due to can affect the sex-ratio of populations, this could be a highly useful
cannibalism was 99.2% during the naupliar stages. Nauplii stages of tool to optimize copepod production. There are many factors to
R. nasutus developed escape behaviors to avoid predation from adults understand in order to optimize the sex ratio of copepod cultures and
(Mullin and Brooks, 1967). Lazzaretto and Salvato (1992) reported increase the productivity and research in this direction will be
the existence of cannibalistic behavior in T. fulvus females towards necessary in the future.
first-stage non-related nauplii, which demonstrates that mothers can
spare their own offspring. Yet, if this mother/offspring recognition 2.6. Selection and cross-breeding to obtain optimized populations
raises interesting questions about chemically mediated communica-
tion between copepods, then it would not matter to intense cultures In all food production sectors, production increases by selecting
where young stages face cannibalism from non-related individuals. populations that have particular traits needed for rendering the
The safe way to counteract this effect is to separate nauplii from the production economically valuable. In the marine environment, this
adults (Brandl, 1972). More developed systems as presented by Payne selection procedure started late as compared to land-based food
and Rippingale (2001a) and Zillioux (1969) are of great importance. sectors but improvement of reared marine organisms is done
Kang and Poulet (2000) and Liang et al. (1994) showed that egg intensively (Duarte et al., 2007). The selective reproduction technique
cannibalism was also important, suggesting that separation of the is only applied to a small percentage of the hundreds of organisms
swimming stages and the eggs is crucial. Sedimentation and regular raised in aquaculture, and therefore there is great room for
harvest of the eggs from a culture is necessary to increase the harvest improvement of reared species which have not yet been selected in
of egg production per unit. This regular harvest also keeps tanks clean any way (FAO, http://www.fao.org/biodiversity).
by removing fecal pellets, exuvia, detritus particles, sedimented algae, Selection of groups of individuals has to be combined with the
etc. (pers.obs.). development of techniques for storage and selection of sperm,
embryos and other products from desirable parents (see Conservation
2.5. Optimization of sex-ratio and shipping section). Within the subphylum of Crustacea, prelimi-
nary studies can be realized by using Artemia as a model because
Another way to improve the productivity of copepod cultures is to many of the factors thought to be responsible for genetic differenti-
control the sex-ratio to optimize the number of females producing ation and speciation in other organisms are observable in these
offspring. The males' main role is to fertilize the females. In several organisms (Barigozzi, 1982; in Eimanifar et al., 2006). For instance,
species, one spermatophore is probably enough to fertilize eggs for studies to produce fast hatching Artemia have already been performed
~ 2–2.5 months (e.g. Acartia clausi, Uye, 1981, Calanus spp. Marshall (Briski et al., 2008) and similar studies could be achieved by using
and Orr, 1955, and Eurytemora affinis, Katona, 1975). In other species relevant copepods.
(e.g. Centropages spp., Temora stylifera, and A. tonsa), females lack Indeed, little work on selection has been conducted for copepods,
seminal receptacles and thus the ability to store sperm except in the but the selection of heritable traits that are valuable for aquaculture is
spermatophores and need to replenish their sperm supply continu- an interesting suggestion. The size-dependency of reproductive
ously (Ohtsuka and Huys, 2001). Several studies have shown that performance in male copepods has been examined in A. tonsa
during experimental time females tend to outlive males (e.g. (Ceballos and Kiørboe, 2010). Ceballos and Kiørboe (2010) showed
Rodriguez-Grãna et al., 2010), although the fecundity of the females that large males of A. tonsa produced larger spermatophores than small
does not seem to be affected due to their ability to save sperm. ones, and that the production of offspring in the female increases with
Behavioral and field observations together with population models the size of the spermatophores she received and thus the size of the
suggest that in some species (e.g. Oithona davisae) population growth male. Size is a heritable trait in copepods (Mclaren, 1976; Mclaren and
is severely limited by fertilization rate and by the shortage of males, Corkett, 1978), and females choosing large males may therefore sire
even when population densities are high. In some species (e.g. Temora larger sons and daughters that likewise sire more offspring (Weath-
longicornis) one male can mate with and fertilize more than one erhead and Robertson, 1979). Selection for large males will therefore
female per day. Therefore decreasing the density of males in some optimize the female's egg production. Most likely, the relative stability
species will be cost efficient in terms of feed expenditure and culture of culture conditions (salinity, temperature, and feed type) is likely to
volume. Based on Fisher's (1930) fundamental law of natural result in a genetic selection in the live prey population. An on-going
selection, the sex-ratio should be 1:1. If one sex is in short supply in study using an A. tonsa population validates this hypothesis showing
a population, then an allele that leads to the production of the rarer that phenotypic variability decreased significantly over the first five
sex will be favored (Fisher, 1930). Only when the sex-ratio is exactly generations while under a constant laboratory environment and
1:1 will the expected success of a male and a female be equal and the staying almost constant thereafter. After isolation from the field, the
population stable. Though the sex-ratio in the natural environment is average body size of A. tonsa remained almost stable over 15
often observed as different from what was expected by Fisher's law, generations under controlled laboratory conditions (Halsband-Lenk
this has been explained by a higher mortality of males compared to et al., submitted for publication). However, this relative stability is
females because of their risky motility pattern and speed when probably not developed as far as it would be necessary for aquaculture
seeking mates (higher risk of predation) (Hirst and Kiørboe, 2002; purposes at this point in time.
Kiørboe, 2006, 2007). Sex changes which are believed to be under Recent common garden experiments revealed that different
strong environmental control (food limitation being the strongest copepod populations of the same species could actually exhibit
factor) may be an important mechanism determining the adult sex different life history traits which could be selected to develop a
ratio (Gusmão and Mckinnon, 2009). The fertility of intersex is particular population for aquaculture purposes (Drillet et al., 2008a,b;
160 G. Drillet et al. / Aquaculture 315 (2011) 155–166

Leandro et al., 2006);although none of these studies tried further to et al., 2003; McEvoy et al., 1998; Næss et al., 1995; Toledo et al., 1999).
breed the different populations for numerous generations. However, In these semi-extensive inland tank systems, isolated populations
cross-breeding allopatric populations can both result in an increased from the natural environment are either caught and used directly or
fitness of the following generations (heterosis or hybrid vigor) or grown in tanks; in which occasional phytoplankton blooms are
decreased fitness of the following generations (outbreeding depres- stimulated by the addition of inorganic nutrients to the water. Semi-
sion or hybrid breakdown) and this is due to the recombination extensive systems require low maintenance and allow production of
processes over the multiple generations (Edmands et al., 2005, and large amounts of copepods, eggs and nauplii. However, due to their
references therein). open nature they are prone to be infected or contamination by other
copepod species, zooplankton and parasites or viruses that either
2.7. Enrichment and use of probiotics infect the copepods or use them as intermediate hosts infecting the
fish larvae (Markowski, 1933).
Enrichment of live prey in aquaculture is a well-known practice Whatever the risks of contamination, recent studies on production
and innovation in this field is necessary. However, copepods are often and biochemical composition of copepod production all year round
used as sole feed when compared to enriched prey because work have shown that outdoor culture and wild copepods were highly
performed in this area is mainly focused on comparisons of valuable for fish production (Sørensen et al., 2007; van der Meeren
biochemical composition and effects upon the predator using various et al., 2008). Thus, improvement and sustainable enhancing proce-
live prey organisms reflecting common husbandry in fish rearing. dures are achievable and should be further optimized by a thorough
Among alternative methods developed to increase the production or research effort.
survival of live prey for aquaculture, beneficial bacteria and microbial
preparations gathered under the generic term “probiotics” are now
currently used. These probiotics can contribute to limiting the 3.2. Temperature–salinity–oxygen–light
establishment of pathogenic bacteria (Rengpipat et al., 1998, 2003).
They can also enhance performances of crustaceans by stimulating Temperature, salinity, dissolved oxygen and light are factors that
immunity of the host species like it was observed with Penaeus greatly influence the success of a copepod culture. Most of these
monodon and Peneaus vannamei (Alavandi et al., 2004; Gullian et al., factors have been considered in environmental studies and
2004), or improving the digestive physiology and the stimulation of therefore plenty of literature information is accessible to culturists
digestive enzymes of Litopenaus stylirostris (Frouël, 2007). Probiotic (Mauchline et al., 1998). The primary need for information with
microorganisms might improve the host physiology by contributing regards to large-scale and dense copepod cultures resides in the
to the host nutrition supply of vitamins (vitamin C, vitamin B12, changing effects of these parameters during intensive cultivation.
retinol and others), proteins, amino acids, and highly unsaturated Temperature and salinity have to be adapted to each population of
fatty acids (Gapasin et al., 1998; Gorospe et al., 1996; Yu et al., 1988), copepods and are often analogous to the conditions the population
and delivering enzymes which stay active in the host intestine. is facing in the natural environment because populations have
Microorganisms could facilitate diet assimilation and decomposition adapted to these particular environmental conditions. To our
of refractory compounds (Hood et al., 1971) and convert all dissolved knowledge the optimal temperature and salinity do not change
nutrients, normally unavailable because of inappropriate molecular during intensification processes, preventing extreme conditions is,
size (Verschuere et al., 1999). All these aspects may improve the most of the time, sufficient to avoid a restraining effect on copepod
development and the survival rate of the crustacean by enhancing the cultures.
diet assimilation. Low oxygen tension in culture tanks is most likely to increase the
However, research on the effects of probiotics on copepods is, to mortality and decrease the fecundity of the copepods. Marcus et al.
our knowledge, limited to a few text lines referring to the use of a (2004) reported that 1.4 ml l− 1 O2 (2 mg l− 1 O2) does not affect the
“commercial probiotic A” in a book chapter (Su et al., 2005) where the mortality but decreases the fertility among A. tonsa. At 0.7 ml l− 1 O2
effect of a mixture containing probiotics increased by two fold the (1 mg l− 1 O2), both fertility and mortality are adversely influenced.
production of a copepod culture. Recently, a study where heat However, within a species, some populations experiencing routinely
inactivated Lactobacillus was added to an A. tonsa culture the natural hypoxia could evolve into a more resistant population as
individual size of the adult females and their egg production suggested by the work of Oppert (2006).
significantly increased (Drillet et al., 2011). Considering the potential Light rhythm is an important factor controlling the physiological
of probiotics for copepods, it seems obvious that more research on the performances of copepods, activating sometimes the production of
subject is needed. diapause stages (Marcus, 1982; Alekseev et al., 2007). This factor
affects egg production and hatching success of Acartia sp. cultures;
3. Part 2: Complementary technologies — technical challenges longer light exposure increased the 48 hour hatching success of the
and perspectives eggs (Camus and Zeng, 2008; Peck et al., 2008). However, cultivation
procedures kept constant over time under particular conditions are
3.1. Copepod cultures likely to select for specific traits. For example, cultures of A. tonsa
from the Danish Technical University (DTU-Aqua) have lost their diel
Vijverberg (1989), in a paper aimed at evaluating vital rates of feeding and egg production rhythm due to excess food availability
copepods and cladocerans, noted that marine copepod culture and absence of predators through 150 generations (Tiselius et al.,
techniques were more advanced than fresh water ones. Zillioux 1995).
(1969) presented one of the very first culture systems for copepods, Light spectra is also a parameter that should be studied because
followed by research groups who improved and intensified the ultraviolet (UV) radiations usually generate harmful effects by
system (Payne and Rippingale, 2001a; Schipp et al., 1999; Sun and lowering egg hatching success and increasing mortality, though
Fleeger, 1995). Their technical descriptions are excellent for the some supra-littoral species are more resistant than open water
aquarium trade but more adaptations and improvements are needed species (Chalker-Scott, 1995; Kane and Pomory, 2001; Karanas et al.,
to reach the huge demand of mass cultivation systems required by the 1979; Saito and Taguchi, 2003). Although UV could potential be used
aquaculture industry. as a manipulator to increase pigmentation, since there seems to be a
Wild or semi-wild copepods are used successfully to raise fish tradeoff between risk of predation (low pigmentation) and risk of UV
larvae (Conceição et al., 1997; Engell-Sørensen et al., 2004; Evjemo damage (High pigmentation) in zoo-plankton (Hansson, 2000). Since
 G. Drillet et al. / Aquaculture 315 (2011) 155–166 161

most fish larvae are visual predators (Hunter, 1981) a feed item with quality by using probes to measure O2, pH, salinity and CO2 and are
“higher” pigmentation and thereby more visible would be preferable. widely used in the aquaculture sector to counteract water quality
fluctuations by adjusting pH, O2, salinity using fixed set points.
3.3. Water quality, recirculation and automation of cultures Stable water quality is a necessity to develop copepod mass culti-
vation systems.
Antibiotic treatments to increase survival, growth, and lifespan of
crustaceans including copepods have occasionally been used (Corkett
and Urry, 1968; Fisher and Nelson, 1978; Pelletier and Chapman, 3.4. Conservation and shipping
1996). However, the use of antibiotics at the industrial scale must be
avoided because of potential contamination of the natural environ- An additional challenge for the enlargement of copepod use in fish
ment by resistant bacteria. Also, exposures to antibiotics could farming and in the aquarium trade is the development of storage
generate toxic effects to crustaceans including copepods (Di Delupis processes for various development stages. These techniques would
et al., 1992; Drillet et al., 2007; Flaherty and Dodson, 2005; Isidori allow the shipping of copepod products between producers and end-
et al., 2005; Wollenberger et al., 2000). users and help producers to ensure back-up of cultures for re-
Since the introduction of governmental regulations on waste inoculations and re-stocking of their systems. Payne and Rippingale
water quality from production plants and due to the need for high (2001b) stored fresh nauplii of Gladioferens imparites for up to 12 days
water quality in intensive systems, the aquaculture industry has at 8 °C (survival 99%) but mortality was almost 100% after 36 days.
developed re-circulated fish farming systems. This is because low One alternative resides in the storage of eggs. Marcus and Murray
water quality leads to poor growth and fertility, increased disease (2001) stored diapause eggs of Centropages hamatus for 17 months
pressure, and mortality (Boyd and Tucker, 1998; Schreck et al., 2001). under anoxia keeping a hatching success N80%. Around 50 copepod
The recirculation of water through a proper system helps at species belonging to Acartiatiidae, Centropagidae, Pontellidae and
controlling factors such as NHx compounds, phosphorus, pH and Temoridae have been described to produce resting eggs as part of
CO2, which greatly affect fish/crustaceans welfare. Indoor copepod their life cycle (Mauchline et al., 1998). However, there are plenty of
cultures are often kept in small volumes where water change is easy complexities regarding the induction of diapause and the cues
to perform but scaling-up the volume of culture renders this breaking diapause are scarcely understood. Some authors propose
impossible. A solution resides in the recirculation technologies that peroxide production due to the oxidation of fatty acids to
which should be adapted since copepods may require different prostaglandins by light inside the embryo may trigger hatching in
rearing conditions compared to fish (see typical recirculation Brachionus sp. (Hagiwara et al., 1995), while other authors working on
components in Losordo et al. (1998) and Fig. 1). Artemia spp. manipulated intra-cellular pH to force hatching during
With biofilters, organic and inorganic substances are removed the refractory period (Drinkwater and Crowe, 1987).
using a biologically active biofilm, which is created when bacteria Alternatively, Drillet et al. (2006a) stored subitaneous eggs of A.
grow on the surfaces of the biofilter element (Lazarova et al., 1994; tonsa for up to a year at low temperatures (2–3 °C), forcing eggs to
Juretschko et al., 1998). The great variety of bacteria in the biofilm switch to a quiescent state. The authors showed that cold storage does
makes this technology adaptable to a wide range of environments not affect the fitness of the next generation of copepods, although
although its success to enhance the water quality depends on the they observed an increased mortality during the ontogenetic
ability to control the processes in the biofilter (Tal et al., 2003; Tseng development (likely due to the lack of reserves transferred from the
and Wu, 2004). An efficient control of suspended solids as food stored eggs into the first nauplii). However, the biochemical profile of
residuals, exoskeletons and feces, is essential for the stability of eggs and nauplii is negatively affected by cold storage (Drillet et al.,
biofilters (Okabe, 1996; Piedrahita, 2003). The particle content affects 2006b; Støttrup et al., 1999) and egg ontogeny affects their capacities
the oxygen level, as less oxygen can be dissolved in water with to be cold stored (Drillet et al., 2007). Additionally, under similar
impurities than in clean water also the efficiency of UV disinfection is storage conditions, different populations of A. tonsa showed totally
decreased by the presence of particles in the water column different capacities of cold storage, showing the importance of the
(Summerfelt and Sharrer, 2004). population origin (Drillet et al., 2008a).
UV radiation is the most common water disinfection method in Previous studies have shown that optimal cold storage conditions
aquaculture industries and is used to reduce the presence of harmful of wild harvested eggs from a Danish estuary were: low temperature
bacteria, parasites and viruses in the water. Chlorination–dechlorination (b5 °C), medium salinity (10–20) and anoxia (Holmstrup et al., 2006).
using chlorine and thiosulfate is another effective method to Eggs from a culture of Acartia tonsa even survived storage at room
disinfect water before batch cultivation (see method in Støttrup temperature in distilled water for weeks keeping a viability of 15–20%
and McEvoy, 2003). The use of ozonation is another way of (Højgaard et al., 2008). Thus, in the marine environment eggs can
decreasing the bacterial development in cultures and was shown survive in the sediment as long as 40–69 years (Marcus et al., 1994;
to be effective at low levels (400–500 mV oxidation–reduction Sichlau et al., 2011). This gives researchers a lot of room for
potential), increasing the growth and survival of the southern rock improvement of storage techniques for copepod eggs. Recently, first
lobster (Jasus edwardsii, Ritar et al., 2006). One of the risks related to steps towards cryo-conservation of eggs were conducted by Ohs et al.
copepod cultures in intensive fish farms isthe pollution of copepod (2009) but, as stated by the authors, much more work has to be
cultures by rotifers which may take over the systems. This was partly achieved in this field.
solved in a recent study where the use of flubendazole as a Limitations to the shipping of copepod eggs are the transfer of
therapeutic agent was used to remove Brachionus plicatilis from parasites, histophageous ciliates and bacteria from one system to
copepod cultures (Steenfeldt and Nielsen, 2010). another. Ciliates can be removed from egg samples by simple fresh
Another issue with copepod cultures is that they demand many water rinsing (personal observation) and chemicals can be used to
man hours for daily routines, such as algal culture maintenance, disinfect properly the eggs as they are known to be much more
copepod culture maintenance, sorting copepods, egg harvest, etc. resistant to various chemicals than nauplii (Næss, 1991a,b; Næss
Automation of these procedures with an automatic control of and Bergh, 1994). For egg carriers, Støttrup and McEvoy (2003)
feeding by dosing pumps, sorting of copepods and egg harvest by proposed to disinfect eggs (Tisbe holothuriae Humes) by adding
size fraction mechanical filters, and the use of an efficient chlorine to the water, whereby the egg sacs detach from the dead
electronic control system is a key to success with large-scale adults, can be harvested, rinsed and used to hatch nauplii under
copepod cultures. Automatic control systems ensure stable water axenic conditions.
162 G. Drillet et al. / Aquaculture 315 (2011) 155–166

4. Part 3: Sales and marketing — commercialization strategy for aspects of copepod production. Molejon and Alvarez-Lajonchere
copepod products (2003) proposed to hold productions in countries with low “qualified
labor” costs because in hatcheries from industrialized countries, for
It has been argued over the years that constructing a successful example, salaries and benefits represent the largest cost-component
commercialization strategy is important to capture the value from of production at 37% (Juarez and Martinez-Cordero, 2004). Alvarez-
research and development activities (in our case Copepod products, Lajonchere and Taylor (2003) took copepod cultures into account in
Fig. 2). However, the economic value and commercialization of their calculation of cost efficiency in the production of juvenile
copepods has, to our knowledge, never been assessed even though common snook, but no details for price of production are presented
patents for fish breeding including the use of copepods do exist (price per amount of produced copepod) as can be found for Artemia
(JP2007044042 and CN1288967). Here, we adapted Teece's concep- production (Lim et al., 2001). In the aquarium trade sector, however, it
tual framework (1988) with the objective to identify the major gaps in is possible to find distributors, retailers and customers of live copepod
our knowledge. The framework lists the main parameters influencing products through internet searches, but access to these products is
the success of the product in focus. Some of these have been discussed limited and prices vary drastically from one retailer to another
earlier and the ones related to management are discussed here. We (Table 2). One of the reasons for this could be that firms producing
consider that it is essential to differentiate between centralized vs. copepods sell their products only in their own geographical areas,
decentralized productions when analyzing the economical and using only the local languages and networks, making national or
commercialization aspects. One should like to keep in mind two international research for copepod products relatively difficult. In our
levels of considerations: compilation, only one retailer proposed its products in more than one
of the major languages (Table 2).
1. The copepod production is held on site at the end-user production Another important limiting factor could be the difficulty of import/
plant (decentralized production), where the copepods are needed export of live products (Distribution) in rather small amounts across
for first feeding and in this case, the price of the copepod products borders due to custom Regulations which differ from country to
is bound to the capital investment for the production system country and small aquarium businesses may not be well prepared for
(culture system and eventually expertise from consultancy) and international export (Adelaide Rhodes, personal communication). In
the cost of production (labor and variable costs = maintenance, our opinion, economic and market studies are lacking because the
electricity, consumables etc.). effects of an improved availability of copepod products could increase
2. The copepod production is completed by a firm that sells copepod the number of fish species raised through their entire life cycle not
products to end-users (centralized production). The copepod only affecting the aquarium trade. An important gain would be, for
producer needs to incorporate in his selling price the price due to example, to preserve natural habitats by decreasing the demand/
the capital investment, cost of production, marketing, and potential
shipping prices. The end-users benefit from a decrease in labor
cost, capital investment cost and need for expertise, but they are Table 2
List of copepod products available from retailers advertising and selling on the internet
also restricted to the price set by the competition/availability of
(January 2009, with authorization from companies). (*1000 = 1000 individuals).
copepod products on the market.
Firm Product price Language
We reviewed and discussed Competitive production and comple- AlgaGen Pseudodiaptomus pelagicus 50 US$ English
mentary Technologies earlier in this paper (Parts 1 and 2) and ask here (*1000)
two simple questions: “How much does a copepod cost?” and “How AlgaGen Acartia tonsa 50 US$ English
(*1000)
does one commercialize a copepod?” The aquaculture sector
AlgaGen Parvocalanus sp. 100 US$ English
occasionally uses copepods from semi-extensive in situ systems but (*1000)
no live copepod products are available for large aquaculture AlgaGen Tisbe sp. 35 US$ English
businesses. Only a few published studies deal with the economic (*1000)
Guernesey Sea Acartia tonsa and Tisbe sp. females 160–255 £ English
Farms (*300 up to *1500) 225–355 €
SeaHorseMania Mix of Calanoid and Cylopoid 29.80 € English
copepods 1000 ml German
French
Spanish
Italian
SeaHorseMania Acartia tonsa adults 65 € English
(*1000) German
French
Spanish
Italian
SeaHorseMania Nitokra lacustris 18 € English
200 ml German
French
Spanish
Italian
SeaHorseMania Acartia tonsa eggs 11 €/18 €/ English
*10,000/*50,000/*100,000/*1,000,000 27.5 €/150 € German
French
Spanish
Italian
Seahorse Gladioferens imparites nauplii AUS $110 English
Sanctuary (*5.000–10.000)
SeaHorseMania Tisbe sp. 200 ml/live ReefPods 15 €/18 € English
Tisbe sp. 200 ml German
French
Fig. 2. ‘How to capture value from a copepod product’. Conceptual scheme modified
Spanish
from Teece (1988) showing the most important parameters influencing commercial-
Italian
ization of copepod products.
 G. Drillet et al. / Aquaculture 315 (2011) 155–166 163

availability ratio of some declining/over-caught fish populations and (e.g. eggs). Copepod cultures of non-native species should be reared
therefore keeping sustainable natural resources. This is even more in closed systems to avoid invasions of surrounding environments.
relevant when taking into consideration that many reef fish are • We recommend feeding dense copepod cultures continuously using
captured using destructive fishing practices such as the use of automatic procedures, rather than feeding once or twice daily to
cyanides generating long lasting or even devastating ecosystem and ensure constant optimal food concentrations in the water.
social impacts (see Barber and Pratt, 1997; Johannes and Riepen, 1995 in • We suggest that water recirculation in copepod cultures is essential
Sadovy and Lau, 2002). In addition, it seems that most of the research for further development because water quality is essential in
and development on copepod cultivation and copepod products is cultures. The recirculation system could be linked to an automatic
performed using public Finance. A systematic study of the price of system for harvest of nauplii, eggs and debris to decrease the
copepod production is necessary to raise the interests of business angels cannibalistic effects and to facilitate the storage of copepod eggs.
and venture capitalists. Also, in association with commercialization, the • We recommend launching research with the purpose of selecting
intellectual property rights (IPRs; i.e. patents, trademarks etc.) must be strains or populations of copepods that can tolerate high densities,
considered because they play a central role behind capturing value from exhibit a fast development and growth with moderate mortality,
knowledge assets (Teece, 1998). Many authors argue that strong IPRs exhibit a relatively high resistance to stress, and present a highly
encourage investment because it gives a fair degree of certainty on qualitative nutritional value for fish larvae
return of investment (but surely it doesn't give an absolute guarantee of • We recommend using light regimes that are not known to increase
profit maximization) though, as stated by Teece (1986): “the protection the production of resting eggs (e.g. 12D/12L and UV free) for
offered by patents is fairly easily enforced”. producing copepods, although this should be adapted for individual
The last important gap in knowledge we identified, though it is a species. Also, we recommend hatching eggs with increased light
rather personal opinion, resides in the fact that apart from a few exposure because this seems to enhance the hatching success of
professionals, too few people in the aquaculture and aquarium some copepod species.
industry actually know about copepods and their potentials as live • We recommend using an automation of copepod cultures to
feed. Screening the internet for copepod businesses, it was obvious decrease labor work and cost of production. We believe that it
that many aquaculture consultants/aquarists do not know enough should be trivial to transfer some technologies from mass fish
about copepods. It is a paradox that knowledge about one of the most farming to copepod mass culturing.
numerous multicellular organisms on the planet and the optimal live • We recommend copepod scientists/culturists/businesses to use/
feed for larval fish exists for researchers but not yet for the obvious develop an efficient and effective Sales and Marketing strategy to
end-users. Hence and to our knowledge, the concept is only utilized in reach and educate consumers and develop a sustainable business.
rare cases. This has to be tightly coupled with a thorough study of economical
feasibility.
5. Conclusions and recommendations

There are several aspects of copepod cultivation techniques and Acknowledgements

use that have been studied and reviewed before (see Lee et al., 2005;
Støttrup, 2003). However many important and novel parameters that We thank all of the copepod culturists, as well as all the copepod
could help the improvement of copepod cultivation techniques businesses who allowed us to publish data from their copepod cultures
should be carefully studied. The scientific community for instance and production. We are also grateful to Jessica Frost for her help with the
should launch research to understand catabolic lipid pathways in language editing of the manuscript and the reviewers of a previous
various copepod species as this would help developing inexpensive version for their valuable comments. This work was supported by GESS
food items for copepods able to bio-convert fatty acids. Also, the need and the Eur-Oecans Network of excellence grants to GD and by the
for understanding mating processes, sex changes, sex-ratio and Danish Agency for Science, Technology and Innovation grants 272-07-
chemical communication between copepods seems obvious and the 0485 to BWH and 10-093759 to GD.
amount of effort put in these areas should be increased before
transferring this knowledge to applied aquaculture. Selection of
interesting life history traits holds great promise in copepod cultures; References
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