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«The impact of dietary fatty acids in common sole larval ( Solea solea, L.) nutrition Lund, Ivar Publication date: Document Version Publisher's PDF, ...»

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The impact of dietary fatty acids in common sole larval ( Solea solea, L.) nutrition

Lund, Ivar

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Lund, I. (2007). The impact of dietary fatty acids in common sole larval ( Solea solea, L.) nutrition. Roskilde

University and Danish Institute for Fisheries Research.

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The impact of dietary fatty acids in common sole larval (Solea solea L.) nutrition PhD thesis by Ivar Lund December 2007 Danish Institute for Fisheries Research Roskilde University Danish Technical University Department of Environmental, Social and Spatial Change The impact of dietary fatty acids in common sole larval (Solea solea L.) nutrition by Ivar Lund Danish Technical University Danish Institute for Fisheries Research Department of Aquaculture The North Sea Centre Roskilde University Department of Environmental, Social and Spatial Change December 2007 Table of contents Preface………………………………………………………………………………………. 4 Dansk resume / Summary…………………………………………………………………… 5

1.0 General introduction……………………………………………………………………

1.1 Objectives……………………………………………………………………………… 10

1.2 Presentation of papers………………………………………………………………..... 11

1.3 List of papers………………………………………………………………………….. 12

2.0 Lipid nutrition in marine fish………………………………………………………….

2.1 Synthesis, desaturation and elongation of fatty acids in fish…………………………... 14

2.2 Eicosanoids…………………………………………………………………. ………… 15

3.0 Broodstock and egg quality………………………………………………..……………

3.1 Wild- and cultured sole breeders; egg viability………………………………………… 17

3.2 Reproduction, - essential fatty acids. Egg and early larval quality……………………. 18

4.0 Live prey species, Tisbe, rotifers and Artemia as feed for sole larvae……….............

4.1 Larval prey capture……………………………………………………………………. 21

4.2 EFA in live prey and dietary EFA requirements in start feeding sole larvae…………. 23

4.3 Transfer of FA between trophic levels and tissue FA interactions……………………. 24

5.0 The influence of essential fatty acids in larval fish respiration………………………

6.0 Pigmentation …………………………………………………………………………… 27

6.1 The inducement of malpigmentation in sole larvae by dietary EFAs…………………. 29

6.2 Larval sensitivity to ARA and pigmentation window……………………………….... 29

6.3 Hormonal influence on pigmentation…………………………………………………. 31

6.4 Mechanisms of PGE2 and pigmentation………………………………………………. 32

7.0 Eye migration……………………………………………………………………………

8.0 Conclusions………………………………………………………………………………

9.0 References……………………………………………………………………………….

Preface The present thesis deals with the importance of fatty acids (FA) in larval nutrition of common sole (Solea solea L.); a species with a high commercial potential in aquaculture. Dietary fatty acids are found very important for the optimal development of most marine fish larvae and this has only been sparsely examined in common sole larvae. A main objective was to characterise the importance and influence of dietary polyunsaturated fatty acids (PUFAs) in relation to quality parameters of sole eggs, larval composition and larval requirements with a specific focus on pigmentation.

The thesis is submitted to Roskilde University, Department of Environmental, Social and Spatial Change, 4000 Roskilde, Denmark in partial fulfilment of the requirements for obtaining the Doctor of Philosophy (PhD) degree. The dissertation consists of 6 manuscripts.





I would like to thank Danish Institute for Fisheries Research for funding this work.

I wish to thank my supervisors Dr. Benni Winding Hansen, Roskilde University and Dr. Svend Jørgen Steenfeldt, Danish Institute for Fisheries Research (DIFRES) for general supervision, support and critical revision of manuscripts and in addition thanks to senior researcher Dr. Erik Hőglund, DIFRES.

A special thank to the laboratory staff at Roskilde University, Department of Environmental, Social and Spatial Change, especially chemist Peter Christensen for assistance with fatty acid analyses and GC-MS equipment. Further thanks to Head of Aquaculture section, Per Bovbjerg Pedersen, Danish Institute for Fisheries Research for general good advice.

Finally I wish to thank my wife, Birgitte for her patience, when my work was first priority… at the expense of the family.

Hirtshals, December 2007Ivar Lund

Dansk resume I bestræbelserne på at afdække mulighederne for opdræt af nye arter i akvakultur er der på Danmarks Fiskeriundersøgelser (DFU i Hirtshals) gennemført et flerårigt projekt med almindelig tunge (Solea solea). For mange marine fiskearter er larvefasen ofte den kritiske periode. Kvaliteten af marine fiskelarver er i stor udstrækning relateret til indholdet af og sammensætningen af fedtsyrer i foderet. Nærværende afhandling omhandler en undersøgelse af indvirkningen af fedtsyrer på kvaliteten af tunge æg samt på tunge larvers overlevelse, vækst, stress respons og pigmentering. Der blev specifikt fokuseret på betydningen af de langkædede flerumættede essentielle n-3 og n-6 fedtsyrer de såkaldte PUFAs.

Det blev påvist at tunge larver har et lavt krav til essentielle fedtsyrer, således var vækst og overlevelse af larver holdt under optimale fysiske betingelser ikke påvirket af forholdet mellem essentielle fedtsyrer eller af fodertyper med lavere indhold af PUFAs. Tunge larvers fedtsyresammensætning afspejlede fødens profil. Forsøgene viste dog, at de essentielle fedtsyrer kan være vigtige for øget stress tolerance under suboptimale forhold som hypoxi.

Den essentielle fedtsyre, arachidonsyre, C20:4n-6 forårsagede dorsal fejlpigmentering (hypomelanosis) som følge af biokemiske reaktioner der involverede hormonstoffer som prostaglandiner. Graden af fejlpigmentering øgedes med stigende koncentration af arachidonsyre og tungelarvers sensitivitet var væsentligt højere forud for metamorfosen. Andre essentielle fedtsyrer som eicosapentaensyre, C20:5n-3 og docosahexaensyre, C22:6n-3 var tilsyneladende ikke involveret i pigmenteringen. Stærkt fejlpigmenterede juvenile tunger havde en lavere vækst end ikke fejlpigmenterede, hvilket kan være forårsaget af processer under metamorfosen, idet disse fisk var kendetegnet ved ufuldendt abnormal øjevandring efter endt metamorfose. Fejlpigmentering på undersiden (overpigmentering, hypermelanosis) var specifikt relateret til enkelte larvehold og påvirket af fysiske forhold.

Summary The thesis focuses on the importance of dietary fatty acids (FA) in relation to the quality of sole eggs, - larval pigmentation development and physiological requirements of sole larvae (Solea solea L.) held under experimental rearing conditions. Sole is a promising candidate for aquaculture, but fatty acid requirements have been sparsely examined. Dietary fatty acids are very important for most marine fish larval performance, especially the long chained polyunsaturated fatty acids, PUFAs. A major problem in flatfish farming is skin malpigmentation, which may be influenced by unbalances in tissue n-3: n-6 PUFA composition. A main objective was therefore to elaborate knowledge on the effect of essential dietary n-3 and n-6 PUFAs on the pigmentation process of sole larvae.

The FA profile of cultured eggs was affected by the pelleted diet of the F1 cultured broodstock and wild or culture origin may be determined based on the egg FA profile. The differences in specific fatty acids profiles, however did not influence subsequent larval growth performance.

Common sole larvae may be start fed on Artemia. Unenriched Artemia strains of various FA qualities proved to be fully as good as essential fatty acid, EFA enriched Artemia with respect to larval growth, survival and pigmentation. Rotifers and the harpacticoid copepod Tisbe holothuriae should not be used in common sole larvi culture as they did not promote a good growth of the larvae or any gains to the culture. This may be related to their smaller size (i.e energy content per prey) more than any fatty acid deficiencies. In relation to the growth performance of common sole larvae highly expensive commercial dietary FA emulsions seemed only marginally better than a mixture of marine fish oil, E vitamin and soy lecithin. Both growth rate and survival of sole were not influenced by the dietary essential fatty acid (EFA) content. Hence, common sole larvae seem to have a relatively low requirement for dietary eicosapentaenoic acid, EPA and docosahexaenoic acid, DHA in comparison to several other flatfish species. However, an increase in stress resistance was observed in sole larvae exposed to hypoxia when fed EFA enriched live prey contrary to unenriched live prey. This is believed to be related to a more optimal dietary content of essential fatty acids.

The FA profile in well growing common sole larvae highly reflected the dietary feed and the profile changed within a relatively few days in favour of a new diet.

Increasing dietary and tissue levels of arachidonic acid, ARA caused a displacement of EPA and DHA content, when these were not supplemented in the diet. This was probably related to a higher cell affinity to ARA. Sole larvae seemed to retain tissue ARA and especially starving sole retained the FA profile quite conservatively.

The degree of malpigmentation, - hypomelanosis in sole larvae fed ARA was dose dependent and highly related to the dietary ARA concentration. However, larval sensitivity to dietary ARA in relation to hypomelanosis was considered much higher during pre metamorphosis than during metamorphosis. Further, the sensitivity of sole larvae to dietary ARA induced hypomelanosis was batch related. Live prey enriched by dietary ARA oil of 10 percent or more induced hypomelanosis with no interactive effects by a concomitantly dietary inclusion of EPA and / or DHA. Blind side pigmentation, - hypermelanosis was induced by environmental factors. Larvae may be predisposed to hypermelanosis, which may explain differences in batch related ARA induced hypomelanosis sensitivity.

The cause of malpigmentation induced by ARA may involve eicosanoids of the type prostaglandins, PGE2, as a positive correlation was observed between an increase in larval tissue PGE2 concentration, ARA tissue content and malpigmentation.

Initiation of metamorphosis defined as initiation of eye migration was related to the body length of the larvae (i.e. when larvae were 7-8 mm) and not to the diet or EFA treatment. The advance of metamorphosis (i.e. eye score) was similarly related to larval length. For the majority of albinic juveniles eye migration was impaired, and resulted in a permanent incomplete aberrant non symmetrical eye position. This abnormality was not solely related to the effect of dietary ARA or some of its metabolites, as it was also observed for albinic fish not treated with dietary ARA. It is regarded as a trait related to albinism and dietary caused albinism may therefore involve morphological changes in sole larvae.

Highly malpigmented or albinic juvenile fish had a slower growth rate than normal pigmented juveniles, which may be related to visual abilities in detecting prey.

1.0 General introduction The last two decades have witnessed a remarkable global increase in the culture of marine finfish and globally aquaculture is probably the fastest growing food producing sector accounting for 33 percent in 2004 increasing to 50 percent of the worlds food fish in 2006 (FAO, 2006).

Figure 1.

Illustration of development of aquaculture production and fisheries (FAO, 2003)

–  –  –

from the northwest African coast and Mediterranean in the south to the Irish Sea, southern North Sea, Skagerrak and Kattegat in the north.

Figure 2. Illustration of geographical distribution of common sole

–  –  –

The present work on Common sole larvae was an extension of this research project, financially supported by a PhD fellowship from Danish Institute for Fisheries Research.

Broodstock and larval nutrition are central research areas within marine aquaculture; as because of limited knowledge within these segments; and due to the increasing need for superior quality fish fry for both the aquaculture industry and restocking. Unpredictable and variable egg quality is a major limiting factor for successful mass production of juveniles and dietary essential fatty acid, EFA content has been found to greatly affect spawning quality in fish (Izquierdo, 1988, FernándezPalacios, et al., 1995).



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