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Echinoderms :San Francisco, Mooi & Tel<strong>for</strong>d (eds) © 1998 Balkema, Rotterdam, ISBN 90 5410 929 7<br />

<strong>Evaluating</strong> <strong>artificial</strong> <strong>diets</strong> <strong>for</strong> <strong>small</strong> <strong>Paracentrotus</strong> <strong>lividus</strong><br />

(Echinodermata: Echinoidea)<br />

Catherine Fernandez,<br />

CEVAREN, EqEL, Université de Corse, & CRITT Corse Technologie Corte, France.<br />

Charles François Boudouresque<br />

UMR DIMAR 3, Faculté de Luminy, Marseille, France.<br />

ABSTRACT: Artificial <strong>diets</strong> are important <strong>for</strong> sea urchin aquaculture. We studied growth and nutrition<br />

parameters of <strong>small</strong> <strong>Paracentrotus</strong> <strong>lividus</strong> fed 3 <strong>artificial</strong> <strong>diets</strong>. Three different diet types were used :<br />

“vegetable-base” type (low protein - high carbohydrate level), “mixed-base” type (animal and vegetable)<br />

(medium protein and carbohydrate level) or “animal-base” type (high protein - low carbohydrate level). We<br />

recorded ingestion, absorption, assimilation efficiency and total growth rate. These parameters were<br />

recorded monthly over a 9 month period from October 1993 to July 1994. At the end of the experiment, we<br />

also recorded gonadic, test, gut and lantern growth rate. Diet type influenced ingestion, absorption,<br />

assimilation efficiency and total growth rates. The highest ingestion rate was obtained with vegetable feed.<br />

The highest absorption rate was recorded with animal feed, the lowest with vegetable feed. Absorption was<br />

negatively correlated with ingestion and carbohydrate level of the food. The assimilation efficiency was<br />

lowest with vegetable diet and highest with animal diet. Growth of test and gonad was best <strong>for</strong> <strong>small</strong> urchins<br />

fed mixed diet.<br />

INTRODUCTION<br />

The world annual consumption of sea urchins has<br />

been steadily increasing over the last years and<br />

many countries, such as France, are currently<br />

confronted with the problem of over- exploitation<br />

of stocks. For this reason, sea urchin aquaculture<br />

seems to be a promising means of solving this<br />

problem in the future (Le Gall & Bucaille, 1987). In<br />

France recent research aims have been directed<br />

towards the study of <strong>Paracentrotus</strong> <strong>lividus</strong><br />

(Lamarck) in order to establish optimal rearing<br />

conditions <strong>for</strong> this species (Le Gall & Bucaille ,<br />

1987; Fernandez & Caltagirone, 1994; Grosjean &<br />

Jangoux, 1994; Fernandez et al., 1995). One of the<br />

essential points <strong>for</strong> the development of sea urchin<br />

aquaculture is the availability of an <strong>artificial</strong> feed.<br />

In nature, <strong>Paracentrotus</strong> <strong>lividus</strong> feeds essentially on<br />

marine plants although animal material may also be<br />

part of its diet (Neill & Pastor, 1973; Verlaque,<br />

1987). In order to maximize the feasibility of sea<br />

urchin farming (availability, stocking, etc.), the<br />

utilization of <strong>artificial</strong> feed seems indispensable.<br />

Artificial feeds are successfully used elsewhere to<br />

rear Echinoidea or maintain them in aquaria<br />

(Lawrence et al., 1989; 1992; Klinger et al., 1994).<br />

It is there<strong>for</strong>e necessary to understand how of<br />

651<br />

<strong>artificial</strong> feeds are utilized by sea urchins. The aim<br />

of this study is to monitor parameters of nutrition in<br />

<strong>small</strong> sized <strong>Paracentrotus</strong> <strong>lividus</strong> fed different<br />

types of <strong>artificial</strong> feeds measuring ingestion,<br />

absorption, assimilation and growth rates.<br />

MATERIALS AND METHODS<br />

Sea urchins, <strong>Paracentrotus</strong> <strong>lividus</strong>, were collected<br />

in the Urbinu lagoon (Corsica, France,<br />

Mediterranean) in September 1993. The size of the<br />

urchins collected was between 20 and 25 mm<br />

(mean size : 23.2 ± 1.1 mm and mean weight : 5.8<br />

± 0.9 g; mean ± standard deviation). Urchins were<br />

maintained in laboratory in three aquaria filled with<br />

running water (ambient temperature and salinity<br />

between 38 and 39 %). Aquaria were divided into<br />

10 equal compartments and one sea urchin was<br />

placed in each compartment. Urchins were fed<br />

<strong>artificial</strong> feeds during a 9 month period (from<br />

October 1st, 1993 to July 1st, 1994). Each group of<br />

10 individuals (each aquarium) was fed a different<br />

<strong>artificial</strong> feed, which differed in quality and<br />

biochemical composition. The first feed consists of<br />

vegetable meal and vegetable oils: this food is rich<br />

in soluble carbohydrates (58%) and is refereed to as


Echinoderms :San Francisco, Mooi & Tel<strong>for</strong>d (eds) © 1998 Balkema, Rotterdam, ISBN 90 5410 929 7<br />

« vegetable feed ». The second feed consists of fish<br />

meal and vegetable meal in equal quantities mixed<br />

with fish oil and vegetable oil and is referred to as<br />

« mixed food ». Its biochemical composition is a<br />

mix of soluble proteins (29%) and soluble<br />

carbohydrates (35%). The third feed consists of fish<br />

meal and fish oil and is rich in soluble proteins (47<br />

%) and is referred to as « animal feed ». Sea urchins<br />

were always provided surplus food.<br />

Ingestion rates were measured monthly. Ingestion<br />

was determined over a three day period and <strong>for</strong><br />

each feed type, after a given amount of food was<br />

provided every 24h to each urchin. The feed not<br />

ingested at the end of 24 hours was collected, freeze<br />

dried and weighed. Individual ingestion rates were<br />

calculated as the difference between the amount of<br />

food available and the amount of food not ingested<br />

(dry weight). In addition, water content and<br />

dissolution rates of 10 blocks of feed were<br />

determinated. These parameters allowed <strong>for</strong> an<br />

assessment of food biomass losses and were used to<br />

correct daily ingestion rates.<br />

Absorption rates were also measured monthly.<br />

Absorption was determinated by collecting fecal<br />

pellets produced by 10 urchins every 24 hours<br />

during the 3 day feeding period. Food remains,<br />

podia and spines were removed from fecal pellets.<br />

Pellets were then rinsed with distilled water, freeze<br />

dried and weighed. Absorption rates were<br />

calculated using the following equation :<br />

Absorption rate (%) = (ingested biomass -<br />

defecated biomass) x 100 / (ingested biomass)<br />

Growth was also recorded monthly by<br />

determinating the total wet weight (after 1 min<br />

drainage on paper towel) of each sea urchin. At the<br />

end of the experiment the gonad, gut, test and<br />

lantern dry weights <strong>for</strong> each sea urchin were<br />

determined after dissection and drying in an oven at<br />

70° C (to a constant weight). These measurements<br />

were also taken at the beginning of the experiment<br />

from ten urchins collected at the same time as the<br />

experimental animals. The difference in mean<br />

weight <strong>for</strong> each sea urchin part at the beginning and<br />

at the end of the experiment was used to estimate<br />

growth rate <strong>for</strong> each organ. From these data the<br />

assimilation rates were calculated using followed<br />

equation:<br />

Assimilation rate (%) = Ingestion rate x 100 /<br />

growth rate<br />

The nutritional budgets, in dry weight, were then<br />

established from all the data obtained. This data is<br />

652<br />

represented in the <strong>for</strong>mat used by Hawkins &<br />

Hartnoll (1983) as modified by Frantzis.<br />

After testing the normality of the data and variance<br />

homogeneity, the results were subjected to one-way<br />

or two-way ANOVA, coupled with the Tukey test<br />

(Zar 1984).<br />

RESULTS<br />

Ingestion<br />

The sea urchins' mean ingestion rates (dry<br />

weight/day) differed significantly depending on the<br />

feed provided and the time of sampling (two way<br />

ANOVA, p


Echinoderms :San Francisco, Mooi & Tel<strong>for</strong>d (eds) © 1998 Balkema, Rotterdam, ISBN 90 5410 929 7<br />

respectively. The higher the protein content, the<br />

greater the absorption efficiency.<br />

%<br />

90<br />

80<br />

70<br />

60<br />

50<br />

40<br />

30<br />

20<br />

10<br />

0<br />

ABSORPTION EFFICIENCY (%)<br />

49%<br />

Vegetable<br />

feed<br />

67%<br />

Mixed<br />

feed<br />

76%<br />

Animal<br />

feed<br />

Figure 2 : Mean absorption efficiency (± confidence<br />

interval) <strong>for</strong> <strong>Paracentrotus</strong> <strong>lividus</strong> fed three food types.<br />

Growth<br />

The initial size of the sea urchins was 23.2 mm<br />

(5.8 g). At the end of the experiment the final size<br />

varied according to the food provided (one-way<br />

ANOVA, p


Test diameter (mm)<br />

34<br />

32<br />

30<br />

28<br />

26<br />

24<br />

22<br />

20<br />

Test<br />

Diameter<br />

Echinoderms :San Francisco, Mooi & Tel<strong>for</strong>d (eds) © 1998 Balkema, Rotterdam, ISBN 90 5410 929 7<br />

Wet weight (g)<br />

17<br />

15<br />

13<br />

11<br />

9<br />

7<br />

5<br />

Wet<br />

Weight<br />

654<br />

Indices (% wet weight)<br />

60<br />

50<br />

40<br />

30<br />

20<br />

10<br />

0<br />

Gonad<br />

index<br />

Gut<br />

index<br />

Begin of experiment Vegetable feed Mixed feed Animal feed<br />

Lantern<br />

index<br />

Figure 3 : Mean test diameter, wet weight and indices (± standard deviation) <strong>for</strong> <strong>Paracentrotus</strong> <strong>lividus</strong> at the begin of<br />

the experiment and at the end of the experiment <strong>for</strong> the three food types.<br />

Size 20-25 mm<br />

Ingested food<br />

{<br />

v : 34.7 g<br />

m : 25.9 g<br />

a : 25.6 g<br />

v : 49.4 %<br />

m : 66.9 %<br />

a : 75.6%<br />

{<br />

Absorption<br />

F<br />

a<br />

e<br />

c<br />

e<br />

s<br />

{ v : 50.6 %<br />

m : 33.1 %<br />

a : 24.4 %<br />

v : 0.1 %<br />

Gut :{ m : 0.2 %<br />

a : 0.2 %<br />

v : 0.7 %<br />

Lanterne :{ m : 1.1 %<br />

a : 1.1 %<br />

L<br />

v : 6.5 %<br />

o<br />

s<br />

Test :{ m : 10.6 %<br />

s<br />

a : 12.3 %<br />

e<br />

s<br />

v : 0.8 %<br />

Gonads :{ m : 2.3 %<br />

a : 1.6 %<br />

v : 41.3 %<br />

m : 52.8 %<br />

a : 60.5 %<br />

{<br />

Figure 5 : Nutritional budgets (in dry weight) obtained <strong>for</strong> <strong>small</strong> <strong>Paracentrotus</strong> <strong>lividus</strong> over a 9 month period. v :<br />

budget obtained with vegetable feed; m : budget obtained with mixed feed, a : budget obtained with animal feed.<br />

Results of the present study reveal that ingested<br />

food is mainly used <strong>for</strong> test and gonadic growth but<br />

that the levels allocated to each of these tissues<br />

differs depending on food composition (two-way<br />

ANOVA, p


Echinoderms :San Francisco, Mooi & Tel<strong>for</strong>d (eds) © 1998 Balkema, Rotterdam, ISBN 90 5410 929 7<br />

rate when the level of soluble proteins in the food is<br />

low (the level of proteins being an indication of<br />

food quality). Other authors have made similar<br />

observations namely sea urchins ingest greater<br />

quantities of low nitrogen food to obtain the protein<br />

necessary <strong>for</strong> their growth and maintenance (Miller<br />

& Mann, 1973; Lowe & Lawrence, 1976). For<br />

seasonal variations of ingestion rate, similar<br />

observations have already been observed <strong>for</strong> other<br />

Echinoidea: Miller & Mann (1973) observed a<br />

significant positive correlation between ingestion<br />

rate and water temperature <strong>for</strong> Strongylocentrotus<br />

droebachiensis and Fuji (1967) observed that<br />

Strongylocentrotus intermedius feeds very little just<br />

prior to and during its spawns.<br />

In the literature, as in our work, absorption<br />

efficiencies measured <strong>for</strong> <strong>Paracentrotus</strong> <strong>lividus</strong><br />

vary according to food type (Lawrence et al. 1989,<br />

Frantzis, 1992). This last author obtained<br />

absorption efficiencies from 26% <strong>for</strong> Corallina<br />

elongata to 96% <strong>for</strong> Asparagopsis armata.<br />

Lawrence et. al. (1989), using <strong>artificial</strong> foods,<br />

obtain values between 8 and 34 %. The differences<br />

observed could be due in part to the feeds<br />

containing vegetable meal which are rich in soluble<br />

carbohydrates also contain insoluble carbohydrates.<br />

These insoluble carbohydrates are not digested by<br />

Echinoidea (Lawrence, 1976) and would there<strong>for</strong>e<br />

lower the absorption efficiency.<br />

Investigations on <strong>for</strong> other species of echinoids,<br />

have provided evidence of the effect of food type<br />

on growth (Lawrence, 1975 ; Vadas, 1977 Larson et<br />

al.,1980). The good efficiency of feeds contained<br />

fish meal in our experiment confirm that P. <strong>lividus</strong><br />

is omnivorous. The efficiency of an omnivorous<br />

diet <strong>for</strong> sea urchin growth is debated. For<br />

Strongylocentrotus droebachiensis, experiments<br />

suggested that this urchin grew faster on a diet of<br />

kelp than on diet of only mussel (Briscoe & Sebens,<br />

1988) but other experiments suggest that mixed diet<br />

(kelp with ectoproct) is more efficiency than kelp<br />

alone (Nestler & Harris, 1994). Our experiment as<br />

these last experiments suggest that the nutrients<br />

available from omnivorous or animal diet allow <strong>for</strong><br />

optimal growth. This is also true <strong>for</strong> gross<br />

assimilation efficiency. Food composition generally<br />

influences gross assimilation efficiency in<br />

Echinoidea (Fuji, 1967; Leighton 1968), the gross<br />

assimilation efficiencies obtained with animal and<br />

mixed feeds are greater than those observed <strong>for</strong> sea<br />

urchins fed a natural marine plant diet. The<br />

possibility exist that assimilation could be<br />

improved <strong>for</strong> <strong>artificial</strong> <strong>diets</strong>.<br />

655<br />

Concerning organ allocation, several experimentations<br />

have shown that resource allocation<br />

varies under different level of nutrition (Ebert,<br />

1996). Generally, when food is scarce, individuals<br />

allocated relatively more of their available<br />

resources to lantern growth (Ebert, 1980; Black et<br />

al., 1982 ; Levitan 1991) and less to gonad and gut<br />

growth (Ebert, 1996). Our research also suggests<br />

that the quality of food affect resource allocation.<br />

When nutritional value is high (rich in protein), sea<br />

urchin allocated relatively more energy to test and<br />

gonad growth than when nutritional value is lower.<br />

In conclusion, our results suggest a mixed<br />

protein/carbohydrate feed provides the best results<br />

with a low ingestion rate and high absorption and<br />

gross assimilation efficiencies. Test and gonad<br />

growth were enhanced with the mixed diet.<br />

There<strong>for</strong>e we recommend this diet <strong>for</strong> sea urchin<br />

aquaculture.<br />

ACKNOWLEDGEMENTS<br />

The authors wish to thank L. Bronzini de Caraffa,<br />

manager of the SCORSA Company, who permitted<br />

us to work in the Urbinu lagoon and who provided<br />

sea transportation and to Dr. J. Le Campion, Station<br />

Marine d'Endoume, <strong>for</strong> his help in the stastical<br />

aspects of this study. Finally, we express our<br />

gratitude to Dr. D. Viale, University of Corsica, <strong>for</strong><br />

providing laboratories facilities.<br />

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