Elsevier

Aquaculture

Volume 194, Issues 1–2, 1 March 2001, Pages 107-121
Aquaculture

Advances in the development of microdiets for gilthead seabream, Sparus aurata: a review

https://doi.org/10.1016/S0044-8486(00)00501-9Get rights and content

Abstract

The performance of microdiets (MDs) for larvae of marine fish is frequently improved when they are co-fed with Artemia. This suggests that nutritional factors in the live food are positively influencing the ingestion, digestion and assimilation of the MD. This paper reviews recent advances in MD development on the gilthead seabream with special emphasis on studies that isolated, identified and tested these live food factors in MD with the aim of improving their performance.

MD ingestion rates in gilthead seabream larvae increased up to 120% when the fish were exposed to the visual and chemical stimuli of various concentrations of Artemia nauplii. The free amino acids (FAA) alanine, glycine and arginine and the compound betaine were identified from the Artemia rearing medium as metabolites, which stimulated this larval response. Similarly, MD supplemented with phospholipids (PL), particularly phosphatidylcholine (PC), stimulated feeding activity and was consumed up to 45% better in young larval seabream. Moreover, dietary PC appears to have in parallel and/or in tandem a postprandial enhancing effect on lipoprotein synthesis, resulting in improved transport of dietary lipids from the mucosa of the digestive tract to the body tissues.

Live food may also contribute exogenous enzymes to the digestion process or provide factors that stimulate larval pancreatic secretions or activate gut zymogens. Seabream larvae ingesting MD supplemented with porcine pancreatic extract (0.05% DW diet) showed a 30% increase in assimilation and demonstrated significantly (P<0.05) improved growth. Older seabream larvae showed 6.75 times more radioactivity in tissue lipids when fed 14C-triacylglycerol (TAG)-labeled MD supplemented with porcine lipase, while younger larvae demonstrated no improved assimilation.

Factors in live Artemia may influence digestion by stimulating an endocrine response. This was shown when Artemia consumed by seabream larvae elicited a 300% increase in the level of the digestive hormone bombesin compared to levels in larvae given only a MD. On the other hand, liposomes containing the FAA methionine ingested by halibut juveniles elicited higher levels of the digestive hormone cholecystokinin (CCK) compared to juveniles ingesting liposomes containing physiological saline or fish extract. These studies suggested that mobilizing the native endocrine factors associated with the feeding and digestive processes could improve MD performance in gilthead seabream and other species by maximizing its utilization.

Introduction

Rotifers (Brachionus plicatilis) and Artemia nauplii are used extensively worldwide as the dominant live food for the larval stages of both freshwater and marine species with a commercial potential. However, the cost in infrastructure, labor and energy to mass culture these zooplankters represent a significant outlay in investment and running costs. Moreover, the provision of live food is characteristically plagued with variable supply and nutritional quality Sorgeloos, 1980, Watanabe et al., 1983. Consequently, a great deal of interest has been generated to develop an off-the-shelf artificial larval microdiet (MD) as an economic live food alternative. Moreover, MD offer the opportunity to introduce nutrients into the larvae that are not available in the live feed (Rosenlund et al., 1997). However, larviculture based on MD has proved to be an elusive goal as solving the dual complexities of the evolving nutrient requirements of the larvae and the technology necessary to provide a suitably sized, attractive and digestible food particle for the larvae, has been daunting.

Although weaning the larvae from Artemia onto an MD can be achieved at metamorphosis (0.5–0.75 g) in many species Dabrowski, 1984, Foscarini, 1988, Hardy, 1989, the early introduction of prepared diets as the sole replacement for live food has met with limited success Adron et al., 1974, Barnabé, 1976, Kanazawa et al., 1982, Appelbaum and Van Damme, 1988, Walford et al., 1991. Early on, workers realized that the poor performance of MD is related to the variable acceptance and attraction of the inert particle for the larvae compounded by inadequate ingestion, digestion and assimilation. Tandler and Kolkovski (1991) found that Sparus aurata larvae consumed an experimental MD at about one tenth the rate of feeding live food (dry weight) which was sufficient as a maintenance ration but could not support good larval growth and survival.

Section snippets

The enigma of co-feeding suggests nutritional factors in the live food are lacking in MD

Despite the poor performance of MDs when used exclusively to rear marine fish larvae, results were markedly improved when a MD was co-fed with live Artemia nauplii Corneillie et al., 1989, Fermin and Bolı̀var, 1991, Marte and Duray, 1991, Tandler and Kolkovski, 1991, Fernández-Dı́az and Yúfera, 1997, Rosenlund et al., 1997. In seabream larvae, there is a general preference for live food over an inert diet when both are offered simultaneously (Fernández-Dı́az et al., 1994). However, an increase

The modes of influence of dietary factors in Artemia on MD ingestion, digestion and assimilation

The above findings suggest three possible modes of influence by Artemia nauplii on the ingestion, digestion and assimilation of MD during co-feeding: (1) the influence of MD on ingestion by stimulating feeding activity through visual and chemical stimuli; (2) the influence of the biochemical composition of the nauplii on larval digestion and assimilation; (3) the influence of dietary factors that influence both feeding activity as well as digestion and assimilation. Kolkovski et al. (1997a)

The influence of factors in live food on larval MD feeding activity

The free amino acids (FAA) alanine, glycine and arginine and the ammonium base compound betaine were identified as chemical stimuli of gilthead seabream larvae from the 14 metabolites found in the Artemia-rearing medium (Kolkovski et al., 1997a). This was concluded by monitoring the effect of the removal of each of the 14 metabolites on MD ingestion rate in 20-day larvae (Fig. 3). Those metabolites absent in the medium and causing a reduction in the MD digestion rate were considered feeding

The influence of dietary phospholipids (PL) on larval MD feeding activity

Studies carried out at the NCM demonstrated that dietary PL, particularly phosphatidylcholine (PC), stimulate early larval feeding activity but did not elicit the same response in older larvae Koven et al., 1993, Koven et al., 1998, Hadas, 1998. This age-dependent effect is likely a function of the ontogeny of the developing digestive tract which characterizes the larvae of marine teleosts. These authors found that larvae fed with PC-supplemented MD demonstrated up to 35% higher (P<0.05)

The age-dependent influence of dietary PL on larval assimilation

The growth-promoting effect of dietary PL, particularly lecithin, for marine fish larvae is well documented Kanazawa et al., 1981, Kanazawa et al., 1983a, Kanazawa et al., 1983b, Kanazawa et al., 1985, Teshima et al., 1987, Geurden et al., 1997. As this benefit may affect the early juvenile stages Poston, 1990a, Poston, 1990b, the appetite stimulating properties of dietary PC cannot entirely explain the influence of this dietary PL and suggests that post prandial physiological influences are

Do exogenous digestive enzymes influence MD digestion and assimilation?

Another mechanism how live food may directly enhance MD digestion and assimilation is by contributing their own enzymes to facilitate the digestion process until the larva's own alimentary systems is fully differentiated and developed Dabrowski and Glogowski, 1977a, Dabrowski and Glogowski, 1977b, Lauff and Hofer, 1984. In support of this, Kolkovski et al. (1993) found in gilthead seabream larvae ingesting MD supplemented with a porcine pancreatic extract (0.05%), a 30% increase in assimilation

Dietary factors in live food influencing digestion by stimulating an endocrine response

It is well documented that nutrients such as FAAs and FFAs entering the mammalian digestive tract stimulate an endocrine response that controls digestion and nutrient assimilation as well as influencing feeding behavior and food intake. Bombesin and cholecystokinin (CCK), which are pituitary neuropeptides, are an integral part of this gastro-entero-pancreatic endocrine system and have also been reported, or their binding sites, in adult fish Batten et al., 1990, Thorndyke and Holmgren, 1990,

Microdiet prototype for gilthead seabream

The inability of MD to stimulate an optimal digestive response may explain why they are generally poorly digested and, in extreme cases, may cause blockages of the digestive tract (Walford et al., 1991). However, a recent study reported that the capacity of gilthead seabream to digest encapsulated diets depended on the thickness and rigidity of the capsule coating (Fernández-Dı́az and Yúfera, 1995). These authors found, during larval feeding, that thin-walled and soft gelatin microcapsules were

Conclusion

The present review contends that it is insufficient to base MD development on simply imitating the proximate composition of the live food. For optimal MD attraction, digestion and assimilation, specific nutrient factors found in live Artemia, which elicit a physiological response in the larvae, should be incorporated in inert feeds. Future MD research should continue to define, isolate and understand the interdependence of those factors in live food that stimulate feeding activity visually and

References (68)

  • S. Kolkovski et al.

    The effects of live food and dietary digestive enzymes on the efficiency of microdiets for seabass (Dicentrarchus labrax) larvae

    Aquaculture

    (1997)
  • S. Kolkovski et al.

    The mode of action of Artemia in enhancing utilization of microdiet by gilthead seabream Sparus aurata larvae

    Aquaculture

    (1997)
  • M. Lauff et al.

    Proteolytic enzymes in fish developed and the importance of dietary enzymes

    Aquaculture

    (1984)
  • T.J. McDonald et al.

    Characterization of gastrin releasing peptide from porcine non-antral gastric tissue

    Biochem. Biophys. Res. Commun.

    (1979)
  • G. Rosenlund et al.

    Co-feeding marine fish larvae with inert and live diets

    Aquaculture

    (1997)
  • M. Salhi et al.

    Effect of different dietary polar lipid levels on gut and liver histological structure of gilthead seabream (Sparus aurata) larvae

    Aquaculture

    (1999)
  • M.C. Sarasquete et al.

    Histology and histochemistry of the development of the digestive system of larval gilthead seabream, Sparus aurata L.

    Aquaculture

    (1995)
  • M. Thorndyke et al.

    Bombesin potentiates the effect of acetylcholine on isolated strips of fish stomach

    Regul. Pept.

    (1990)
  • J. Walford et al.

    Replacing live food with microencapsulated diets in the rearing of seabass (Lates calcarifer) larvae: do the larvae ingest and digest protein-membrane microcapsules?

    Aquaculture

    (1991)
  • T. Watanabe et al.

    Nutritional values of live organisms used in Japan for mass propagation of fish: a review

    Aquaculture

    (1983)
  • M. Yúfera et al.

    A highly efficient microencapsulated food for rearing early larvae of marine fish

    Aquaculture

    (1999)
  • J.W. Adron et al.

    Rearing of plaice (Pleuronectes platessa) larvae to metamorphosis using an artificial diet

    Fish. Bull.

    (1974)
  • S. Appelbaum et al.

    The feasibility of using exclusively dry diet for rearing of Israeli Clarias gariepinus (Burchell) larvae and fry

    J. Appl. Ichthyol.

    (1988)
  • T.F.C. Batten et al.

    Comparative distribution of neuropeptide-immunoreactive systems in the brain of the greenmolly, Poecillia latipinna

    J. Comp. Neurol.

    (1990)
  • C.L. Cahu et al.

    Early weaning of seabass: are digestive enzymes limiting?

  • C.B. Chan et al.

    Effect of somatostatin on intragastric pressure and smooth muscle contactility of the rainbow trout walbaum

    J. Fish Biol.

    (1992)
  • S. Corneillie et al.

    Reduction of the use of Artemia nauplii by early feeding of sea bream larvae (Sparus aurata) with dry food

  • K. Dabrowski

    The feeding of fish larvae: present “state of the art” and perspectives

    Reprod., Nutr., Dev.

    (1984)
  • K. Dabrowski et al.

    Studies on the proteolytic enzymes of invertebrates constituting the food

    Hydrobiologia

    (1977)
  • K. Dabrowski et al.

    Studies on the role of exogenous proteolytic enzymes in digestive processes in fish

    Hydrobiologia

    (1977)
  • C.H. Dempsey

    Chemical stimuli as a factor in feeding and intraspecific behaviour of herring larvae (Clupea harengus)

    J. Mar. Biol. Assoc. U. K.

    (1978)
  • R.D. Fernald

    Vision

  • C. Fernández-Dı́az et al.

    Feeding behaviour and prey size selection of gilthead seabream, Sparus aurata, larvae fed on inert and live food

    Mar. Biol.

    (1994)
  • I. Geurden et al.

    Effect of a dietary phospholipid supplementation on growth and fatty acid composition of European sea bass (Dicentrarchus labrax L.) and turbot (Scophthalmus maximus L.) juveniles from weaning onwards

    Fish Physiol. Biochem.

    (1997)
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