Protein and Energy Digestibility and Gonad Development of the European Sea Urchin Paracentrotus Lividus (Lamarck) Fed Algal and Prepared Diets During Spring and Fall
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UC Davis UC Davis Previously Published Works Title Protein and energy digestibility and gonad development of the European sea urchin Paracentrotus lividus (Lamarck) fed algal and prepared diets during spring and fall Permalink https://escholarship.org/uc/item/2q5151zv Journal Aquaculture Research, 36 Author Schlosser, Susan Publication Date 2005 Peer reviewed eScholarship.org Powered by the California Digital Library University of California Aquaculture Research, 2005, 36, 972^982 doi:10.1111/j.1365-2109.2005.01306.x Protein and energy digestibility and gonad development of the European sea urchin Paracentrotus lividus (Lamarck) fed algal and prepared diets during spring and fall Susan C Schlosser1, Ingrid Lupatsch2, John M Lawrence3, Addison L Lawrence4 & Muki Shpigel2 1California Sea Grant Extension Program, Eureka, CA, USA 2National Center for Mariculture, Israel Oceanographic and Limnological Research, Eilat, Israel 3Department of Biology,University of South Florida,Tampa, FL, USA 4Texas A&M Shrimp Mariculture Project, Port Aransas,TX, USA Correspondence: S C Schlosser, California Sea Grant Extension Program, 2 Commercial St., Ste. 4, Eureka, CA 95501, USA. E-mail: [email protected] Abstract from the higher dietary energy content of the pre- pared diet. Protein and energy are two of the main limiting fac- tors for sea urchin growth. However, the requirement Key words: Paracentrotus lividus, sea urchin, of daily protein and energy to maximize gonadal pro- gonad, protein, energy,diet duction is still unknown. Paracentrotus lividus were fed three experimental diets: Ulva lactuca, Gracilaria conferta and a prepared diet for 2 months in the fall of 1999 and spring of 2000. Sea urchins from a Introduction laboratory-cultured population of equal age, weight and test diameter were used. Apparent digestibility Aquaculture of sea urchins requires understanding coe⁄cients (ADC%) for protein and energy, using the quantity and quality of algal and prepared feeds acid-insoluble ash as a marker, were measured for for successful gonad production. Paracentrotus lividus all experimental diets. Apparent digestibility coe⁄- (Lamarck) is a commercially important species in the cients for protein was high (475%) for all diets. En- Mediterranean region (Boudouresque & Verlaque ergy digestibility varied among the diets and was 2001). Many aspects of P. l i v i d u s nutrition have been lowest for G. conferta (50^62%). The three diets con- studied including feed type (Fernandez & Boudour- tained varying digestible protein (DP) to digestible esque 1997; Ferenandez & Pergent 1998); comparison energy (DE) ratios of 25,26 and12 mg kJ À1 for U. lac- of di¡erent algal diets (Frantzis & Gre¤ mare 1992) and tuca, G. conferta and the prepared diet respectively. e¡ects of prepared diets (Fernandez & Boudouresque Digestible protein intake was similar for all treat- 2000; Spirlet, Grosjean & Jangoux 2001). The advan- ments, but DE intake was greater for sea urchins fed tage of prepared diets over natural algal diets on sea the prepared diet in both seasons. As a result, the go- urchin gonad growth is known for some species nad production was signi¢cantly higher for urchins (Lawrence, Olave, Otaiza, Lawrence & Bustos 1997; fed the prepared diet, suggesting that energy was lim- Barker, Keogh, Lawrence & Lawrence 1998; Cook, iting in the algal diets. Paracentrotus lividus spawned Kelly & McKenzie 1998) but has not been tested for during the spring experiment, resulting in protein P. l i v i d u s. The cause of greater gonad growth with loss in all treatments. Protein loss was lowest in the prepared diets is not clear. sea urchins fed the prepared diet. Enhanced gonadal Protein, as a main factor in sustaining gonadal growth and gamete development of P. l i v i d u s resulted growth has been examined.Varying dietary content 972 r 2005 Blackwell Publishing Ltd Aquaculture Research, 2005, 36, 972^982 Protein and energy e¡ects on sea urchin gonad development SCSchlosseret al. and using plant or animal protein sources a¡ects bio- ments day À1) in the fall and 21.4 Æ 0.7 1C(n 5120) chemical composition of gonads (Fernandez 1997) in the spring. Dissolved oxygen was 90^100% satura- and gonad production (Lawrence, Fenaux, Corre & tion and mean pH was 8.17 Æ 0.05 (n 512 per experi- Lawrence1991). However, little is known about the ef- ment, one sample week À1 for dissolved oxygen, pH fect of dietary energy and especially the balance be- and salinity). Salinity was constant at 41ppt.The sea- tween protein and energy in the feeds. Most sea water system supplied a continuous £ow of seawater urchin prepared diets contain 20^40% protein and ¢ltered to100 mm to the experimental aquaria. Aqua- digestibility is generally greater than 60% (Frantzis ria dimensions were 20 Â 35 Â 15cm. Each aqua- &Gre¤ mare 1992; Klinger, Lawrence & Lawrence rium was vigorously aerated and contained 10.5 L of 1998; McBride, Lawrence, Lawrence & Mulligan seawater. Sea urchins were always in close proximity 1998; Akiyama, Unuma & Yamamoto 2001), but in- to their food and were checked every morning and formation about energy content or digestibility is evening when temperatures were recorded. generally lacking. The total energy or protein of an Adult P. l i v i d u s were selected from a cohort pro- algal or prepared diet may not represent the quantity duced by spawning individuals in the laboratory in of these nutrients that are available to P. l i v i d u s.Asin December 1997. Animals were starved for 1 week ecological studies, digestibility values of protein and prior to starting both studies to ensure similar nutri- energy of the food consumed are necessary to deter- tional condition for each individual. Sea urchins mine the amount of each nutrient utilized by were measured (horizontal test diameter, HD) to P. l i v i d u s (McClintock1986). the nearest 0.01mm, weighed (whole wet weight, Gonads of sea urchins vary in size and gameto- Æ 0.01g) and placed in replicate, randomly genic state during the year. Gonad production is arranged, glass aquaria (n 5 3 aquaria diet À1, greater in the post-spawning season than in the 10 u rc h i n s a qua r ia À1). Initial mean test diameter spawning season (Lozano, Galera, Lopez, Turon, and whole animal wet weight were 30.7 Æ 1.5 mm Palacin & Morera 1995; Unuma, Kooichi, Furuita, and 13.1 Æ 1.5 g in the fall and 34.3 Æ 1.4 mm and Yamamoto & Akiyama 1996; Lawrence et al.1997; 18.7 Æ 1.7 g in the spring. Klinger et al. 1998). Protein and energy are allocated An initial sample (n 5 20) of P. l i v i d u s, and all ani- to increases in body size or to gonadal production, mals at the conclusion of both studies, were dis- depending on the animals’ reproductive condition sected.Wet body compartments (gonad, gut, lantern, (Edwards & Ebert 1991; Pearse & Cameron 1991; test) were weighed to the nearest 0.01g. The body Fernandez & Boudouresque 2000). compartments were dried at 105 1C for 24 h and re- The purpose of this study is to compare the e¡ects weighed. Interstitial water and coelmic £uid lost dur- of protein and energy on gonad production and body ing dissection and desiccation is not relevant to composition in adult P. l i v i d u s fed algal and prepared protein and energy intake, e⁄ciency or gonad pro- diets during and after the spawning season. Applica- duction. Whole animal dry weight in this study is tion of this information will help improve and opti- the sum of the dry body compartments. Gonad index mize diet formulations for the culture of sea urchins. (%) was calculated as (dry gonad weight (g)/dry body weight (g) Â 100) and as (wet gonad weight (g)/ whole animal weight (g) Â 100). Dry body compart- Materials and methods ment indices are given for gut, lantern and test. One gonad section from each animal in the initial Two identical,60-day studies were conducted during sample (n 5 20) and a sub-sample from each treat- fall (October^December 1999) and spring (March^ ment (n 510 per diet treatment, ¢ve females, ¢ve May 2000). All methods apply to both studies unless males) were preserved in neutral-bu¡ered formalin otherwise stated. All reported values are mean Æ SD. to determine the reproductive state according to By- rne (1990).These were: (1)recovering stage with small previtellongenic oocytes or primary spermatocytes; Experimental conditions (2) growing stage with many large nutritive phago- Seawater £ow from a common manifold was cytes; (3) mature stage with no or few nutritive pha- 0.5 L min À1 to each aquarium with temperature re- gocytes around oocytes or spermatozoa; (4) partly corded twice daily.Dissolved oxygen, pH and salinity spawned stage with some void spaces containing few were measured weekly. Mean seawater tempera- nutritive phagocytes and loosely packed gametes and ture was 24.2 Æ 0.9 1C(n 5120, two measure- (5) spent stage with gonads appearing empty. r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 972 ^ 982 973 Protein and energy e¡ects on sea urchin gonad development S C Schlosser et al. Aquaculture Research, 2005, 36, 972^982 Feeding study Table 1 Composition of algae and prepared diet fed to Para- centrotus lividus in the fall and spring experiments (perg dry The diets fed P. l i v i d u s were fresh, cultured Ulva lactu- matter) ca and Gracilaria conferta (Cohen & Neori 1991) and an extruded moist pellet, hereafter referred to as the Ulva Gracilaria Prepared prepared diet. The prepared diet was manufactured lactuca conferta diet by Wenger International (Kansas, MO, USA).