Effect of dietary phospholipids levels and sources on growth performance, fatty acid composition of the juvenile swimming crab, Portunus trituberculatus

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Effect of dietary phospholipids levels and sources on growth performance, fatty acid composition of the juvenile swimming crab, Portunus trituberculatus
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  Effect of dietary phospholipids levels and sources on growthperformance, fatty acid composition of the juvenile swimming crab, Portunus trituberculatus Xinyu Li a , Jiteng Wang a, ⁎ , Tao Han a , Shuixin Hu a , Yudong Jiang a , Chunlin Wang b a Department of Aquaculture, Zhejiang Ocean University, Zhoushan 316000, China b College of Life Science and Biotechnology, Ningbo University, Ningbo 315211, China a b s t r a c ta r t i c l e i n f o  Article history: Received 11 December 2013Received in revised form 21 March 2014Accepted 26 March 2014Available online 4 April 2014 Keywords: Swimming crab  (Portunus trituberculatus) PhospholipidsGrowth performanceFatty acid An8-weekfeedingtrialwasconductedtoinvestigatetheeffectofvariouslevelsandsourcesofdietaryphospholipids(PL)ongrowth performance and fatty acid compositionof juvenile Portunus trituberculatus . Six semi-puri 󿬁 ed dietswereformulatedtocontainonecontrolgroup,threelevelsofsoybeanlecithin(SL1%,SL2%, SL4%)and twolevelsof eggyolklecithin(EL1%,EL2%),butremainingisolipidticandisonitrogenous.Eachdietwasfedintriplicates(18crabsreplicate − 1 , initially weighing 22.22 ±0.08 g).Thedietary PLsupplementation groups(exceptthe group ofSL 2%)had signi 󿬁 cantly higher weight gain (WG), special growth ratio (SGR) and 󿬁 nal body wet weight (FBW) than thecontrolgroup( P  b  0.05).Moreover,crabsfeddietswith2%ELsupplementationshowedhigherSGRthan2%SLsup-plemented group ( P   b  0.05). With PL supplementation, we also observed that the highly unsaturated fatty acids(HUFA) percentage of the hepatopancreas and muscle increased with a decrease in the polyunsaturated fatty acid(PUFA) level. Compared with the control group, crabs fed diets with EL supplementation had higher n-3/n-6 ratioin muscle ( P   b  0.05). Compared with hepatopancreas, higher level of HUFA (especially for C20:5n-3) and lowerPUFAlevelwereobservedinmuscle.Basedongrowthperformance,thisstudysuggestedthat1%PL(SLorEL)sup-plementedindietscouldsatisfytherequirementofjuvenile P.trituberculatus .Moreover,crabsfeddietswithELsup-plementation had a higher nutritional value of fatty acid pro 󿬁 les than SL groups.© 2014 Elsevier B.V. All rights reserved. 1. Introduction Phospholipids (PL) were evidenced to be an important nutrient forcrustaceans.Themajorityofcrustaceanscouldsynthesizephospholipids,but the level of synthesis is so low that it could not meet metabolic re-quirements (Kanazawa, 1985). Moreover, phospholipids are importantprecursors for a seriesofhighlybiologically activemediators of metabo-lism and physiology, such as diacylglycerol, eicosanoids, etc. (Tocheret al., 2008). As an emulsi 󿬁 er, phospholipids have been reported to im-prove lipidabsorptioninbody(Coutteauetal.,1997).TherequirementsfordietaryPLhavebeenextensivelyconductedincrustaceans,including Litopenaeus vannamei  (Gong et al., 2000; Roy et al., 2006),  Penaeusmerguiensis  (Thongrod and Boonyaratpalin, 1998), and  Eriocheir sinensis (Wu et al., 2010, 2011).It is also well known that phospholipids had a signi 󿬁 cant effect onsurvival,growth,andresistancetostressinsome 󿬁 shesandcrustaceans(Gongetal.,2000;Pascual,1986;Royetal.,2006;Uyanetal.,2007;Wuet al., 2007, 2010, 2011). Due to thedifferent srcins, there were differ-ent fatty acid composition and oxidative stability between egg yolklecithin (EL) and soybean lecithin (SL) (Miyashita et al., 1994; Naraet al., 1997). Several papers have also proved that different PL sources(especially EL and SL) have a different effect on  󿬁 shes and crustaceans(Azarmetal.,2013;Coutteauetal.,2000;Hamzaetal.,2012).Soitisdif- 󿬁 cult to determinethe optimal levels/sources of dietary PL for differentcrustaceans.The swimming crab  (Portunus trituberculatus)  is distributed fromChina, through Korea to Japan (Miyake, 1983). However, because of theover 󿬁 shing and pollutions, the natural resource of   P. trituberculatus  hasshown a downward trend in East China Sea since the 1990s (Yu et al.,2003). Over the past several decades,  P. trituberculatus  has become animportant species for china aquaculture (Chen et al., 2006). However,nutrition studies for juveniles  P. trituberculatus  are still lacking. More-over, the role and requirement of PL in  P. trituberculatus  nutrition havenot been addressed. The objective of the present study was to evaluatethe effect of various levels and sources of dietary PL on growth perfor-mance and fatty acid composition of juvenile  P. trituberculatus , whichmay provide information on the practical use of PL in diets. 2. Materials and methods  2.1. Experimental diets Six semi-puri 󿬁 ed diets were formulated to contain one controlgroup, three levels of soybean lecithin (SL1%, SL2%, SL4%) and two Aquaculture 430 (2014) 166 – 172 ⁎  Corresponding author.http://dx.doi.org/10.1016/j.aquaculture.2014.03.0370044-8486/© 2014 Elsevier B.V. All rights reserved. Contents lists available at ScienceDirect Aquaculture  journal homepage: www.elsevier.com/locate/aqua-online  levels of egg yolk lecithin (EL1%, EL2%), but remaining isolipidtic andisonitrogenous (Table1).ThesourceofSL isalightyellowpowdercon-taining 95% acetone insolubility and 50% phosphatidylcholine (PC)(Siwei Company, Zhengzhou, China). The source of EL is a light yellowpowder containing 90% acetone insolubility and 30% phosphatidylcho-line (PC) (Huaxiahoude Biotechnology Company, Beijing, China). Thefatty acid compositions of the six experiment diets were given inTable 2. During the diet preparation, the  󿬁 sh meal was pulverized andsieved through a 250  μ  m mesh before being added. All dry ingredientswere mixed thoroughly. Fish oil, soybean oil and phospholipids wereadded to the dry mixture and mixed again with water. The diets wereextruded through two types of ori 󿬁 ce diets: 1.5 mm and 2.5 mm.All diets were dried overnight at 50 °C using a hot air oven, and storedat − 20 °C until use.  2.2. Source of crabs and experimental procedures An 8-week growth trial was conducted at the Zhejiang Ocean Uni-versity. The juvenile  P. trituberculatus  were obtained from the Marineand Fishery Research Institute of Zhejiang Ocean University. Therewere three replicates (18 crabs per replicate) for each diet treatment.All crabs were kept separately by using a wheel system (Patent No.201010239818), which was divided in several compartments. Threewheels per tank were used as one replicate. At the beginning of theexperiment, the crabs were acclimated to theindoor cultureconditionsfor1weekwhentheywerefedPL0diet.Then,thecrabswererandomlyselected and stocked into 18 containers of every tank, and fed selectivedietstwicedailyat8:00and16:30.Thedailyrationwasat6 – 8%ofbodyweightduringtheexperiment.Allcrabswereweighedeverytwoweeksandtheirrationadjustedaccordingly.Uneatendietwascollectedtode-termine feed intake. Mortalities and molts were checked and recordedevery day. A photoperiod of 12 h light/12 h dark was maintainedthroughout the experiment. The pH (7.5 – 8.0) and dissolved oxygen(DO  N 6 mg l − 1 ) were monitored regularly. The water temperaturewas 26.7 ± 1.5 °C throughout the trial.  2.3. Sample collection At the end of the feeding trial, crabs were starved for 24 h. All crabsthat survived were counted. The wet weights of the crabs were mea-sured  󿬁 rst. In each treatment, hemolymph samples from four crabswere taken immediately using the method described by Li et al.(2013). All samples were placed in 4 °C for 4 h,followed bycentrifuga-tion(4000rpm,10min, − 4 °C).Thesupernatantwasstoredat − 80 °Cuntil analyzed. Hepatopancreas and muscle from three crabs were dis-sected and weighed to determine the hepatosomatic index (HSI).Then, all samples were frozen immediately and stored at − 80 °C untilneeded.  2.4. Biochemical analysis Moisture, ash, crude protein, and lipid contents of all sampleswere analyzed using standard methods (AOAC, 1995). Crude protein( N   × 6.25) was determined by an auto Kjeldahl System (K358/K355,BUCHI, Flawil, Switzerland). Crude lipid was determined by the ether-extraction method using a Soxtec System HT (E-816, BUCHI, Flawil,Switzerland). Moisture was determined in an oven at 105 °C for 24 h.Ashwasdeterminedusingamuf  󿬂 efurnaceat550 °Cfor12h.Grossen-ergy was determined with an adiabatic bomb calorimeter (HER-15E,Shanghai shangli, Shanghai, China).The activity of superoxide dismutase (SOD) was determined by thexanthine oxidase method described by Lin et al. (2011) and Wang and Chen (2005). The optical density was measured at 550 nm. One unitof SOD activity was de 󿬁 ned as the amount required to inhibit the rateofxanthinereductionby50%ina1-mlreactionsystem.Speci 󿬁 cactivitywas expressed as SOD unit per milliliter hemolymph. The glucose wasdetermined by the glucoseoxidase – peroxidase method (Barham andTrinder, 1972), and the optical density was measured at 505 nm. Totaltriglyceride was determined by the enzymatic colorimetric method ac-cording to Bucolo and David (1973) and Fossati and Prencipe (1982), and the optical density was measured at 546 nm. Total cholesterol  Table 1 Composition of six experimental diets formulated for the  Portunus trituberculatus. Ingredients (%) Experiment diets (phospholipids %)PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%)Fish meal a 60.0 60.0 60.0 60.0 60.0 60.0Soybean oil 4.0 3.0 2.0 0.0 3.0 2.0Soyabean lecithin b 0.0 1.0 2.0 4.0 0.0 0.0Egg yolk lecithin c 0.0 0.0 0.0 0.0 1.0 2.0Fish oil 1.0 1.0 1.0 1.0 1.0 1.0Cellulose 2.3 2.3 2.3 2.3 2.3 2.3Corn starch 19.6 19.6 19.6 19.6 19.6 19.6Monocalcium phosphate 0.5 0.5 0.5 0.5 0.5 0.5Mineral mix d 3.0 3.0 3.0 3.0 3.0 3.0Vitamin mix e 4.0 4.0 4.0 4.0 4.0 4.0Taurine 1.0 1.0 1.0 1.0 1.0 1.0Cholesterol 0.6 0.6 0.6 0.6 0.6 0.6Choline 1.0 1.0 1.0 1.0 1.0 1.0Sodium alginate 3.0 3.0 3.0 3.0 3.0 3.0 Proximate composition (dry matter %) Moisture 8.61 6.99 8.65 5.38 4.98 5.70Crude protein 46.77 46.39 45.73 46.86 45.18 45.82Crude fat 11.48 11.25 10.98 10.37 10.20 10.28Ash 10.16 10.38 10.30 10.51 10.45 10.10Gross energy (KJ/g) 19.61 19.65 19.55 19.34 19.55 19.27 a Purchased from American seafood company, Memphis, USA. b Purchased from Zhengzhou Siwei Company, Zhengzhou, China. c Purchased from Beijing Huaxiahoude Biotechnology Company, Beijing, China. d Mineralpremix,g/kgmixture:calciumdihydrogenphosphate,122.87;lactate,474.22;sodiumdihydrogenphosphate,42.03;potassiumpersulfate,163.83;ferroussulfate,10.78;ironcitrate, 38.26; magnesium sulfate, 44.19; zinc sulfate, 4.74; manganese sulfate, 0.33; copper sulfate, 0.22; cobalt chloride, 0.43; iodate, 0.02; sodium chloride, 32.33; potassium chloride,65.75. e Vitaminpremix,g/kgmixture:thiamineB1,5.0;ribo 󿬂 avin,8.0;nicotinamide,26.0;biotin,1.0;calciumpantothenate,15.0;vitaminB6,3.0;folicacid,vitaminB,5.0;vitaminC,121.0;vitamin K, 2.02; p-aminobenzoic acid, 3.0; vitamin B12, 1.0; cellulose, 504.0; vitamin A, 25.0; vitamin D3, 25.0; vitamin E, 50.0; inositol, 181.0.167  X. Li et al. / Aquaculture 430 (2014) 166  – 172  was determined by the enzymatic colorimetric method according toTrinder (1969), and the optical density was measured at 546 nm. Allparameters were determined by commercial kits (Nanjing JianchenBioengineering Institute, Nanjing, China) using a spectrophotometer(UV-2550, Shimadzu, Kyoto, Japan).  2.5. Fatty acid analysis The muscle and hepatopancreas samples were freeze-dried beforeanalyses (LL1500, Thermo, USA). The lipid in samples was extractedwith chloroform/methanol (2:1  v/v ) according to the method of  Folchet al. (1957). The fatty acid methyl esters (FAME) were analyzed on agas chromatography (GCMS-QP2010, Shimadzu, Kyoto, Japan)  󿬁 ttedwith a SPB-50 (60 m × 0.25 mm id., 0.25  μ  m  󿬁 lm) capillary column(Supelco, Bellefonte, PA, USA). Helium was used as a carrier gas at a 󿬂 ow rate of 44.1 ml min − 1 . The temperatures of the injector anddetector were 200 °C and 250 °C respectively, and the split ratio was50:1. The oven program was 150 °C for 3.5 min, 150 – 200 °C at a rateof 20 °C min − 1 , 200 – 280 °C at a rate of 5 °C min − 1 , and 280 °C for20 min. Results were expressed as the percentage of each fatty acidwith respect to total fatty acids. Fatty acids were identi 󿬁 ed by compar-ing their retention times to authentic standard fatty acid standards(Alltech Co., USA).  2.6. Statistical analysis Data were analyzed using one-way ANOVA, expressed as means ±standard deviation of replicates by Duncan's multiple range test. P   b  0.05 was regarded as statistically signi 󿬁 cant. All statistics wereperformed using the SPSS package (version 17.0). 3. Results  3.1. Survival and growth performance The experiment showed that the test diets were well accepted bycrabs. The survival of crabs from six treatments ranged from 67% to76%, and no signi 󿬁 cant differences were observed among treatments( P   N  0.05). Crabs fed control diet had signi 󿬁 cantly lower weight gain(WG), special growth ratio (SGR) and  󿬁 nal body wet weight (FBW)than other groups ( P   b  0.05), except the group of PL2 (Table 3). TheSGR values of 2% EL supplemented groups were signi 󿬁 cantly higherthan 2% SL supplemented groups and the control group ( P   b  0.05). Inthis trial, feed conversion ratio (FCR), hepatosomatic index (HIS) andmoltingfrequency(MF)ofeachtreatmentshowednosigni 󿬁 cantdiffer-ence ( P   N  0.05).  3.2. Hemolymph biochemical composition Thehemolymphbiochemicalcompositionofjuvenile P.trituberculatus was shown in Table 4. Compared with the control group, diet with PL supplementation increased the triglyceride level slightly ( P   N  0.05).The cholesterol level also increased with EL supplementation, rangedfrom 3.98 ± 1.42 to 6.28 ± 1.39 mg/dl. On the other hand, the choles-terol level showed a tendency to decrease as the SL supplementationlevel increased, and was signi 󿬁 cantly lower in PL3 than PL1 ( P   b  0.05).The glucose level ranged from 2.98 ± 1.12 (PL4) to 4.09 ± 0.86 mg/dl(PL2), but no signi 󿬁 cant differences were observed ( P   N  0.05). Simi-larly, the SOD level showed no signi 󿬁 cant difference among groups( P   N  0.05).  3.3. Body composition Proximatecompositionanalysesofthehepatopancreas,wholebodyand muscle were presented in Table 5. Analysis of the hepatopancreasand whole body samples demonstrated that the dry matter was notsigni 󿬁 cantly affected by PL supplementation ( P   N  0.05). However,the muscle lipid level was signi 󿬁 cantly affected by PL supplementation( P   b  0.05), ranging from 1.00 ± 0.15 (PL3) to 1.46 ± 0.19 (PL2).  3.4. Fatty acid pro  󿬁 le The fatty acid composition (% total fatty acids) of muscle was pre-sented in Table 6. The concentration of  ∑ PUFA in muscle decreasedasPLsupplementationincreased.Moreover,crabfed2%ELdiethadsig-ni 󿬁 cantly lower ∑ PUFA content than other groups ( P   b  0.05). TheC18:1n-9 and C20:5n-3 (EPA) in muscle of juvenile  P. trituberculatus had no signi 󿬁 cant difference among the different groups ( P   N  0.05).  Table 2 Selected fatty acid composition of the experimental diets (% of total fatty acids).Experiment diets (phospholipids %)Fatty acid PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%)C14:0 1.82 1.90 2.07 2.44 1.96 2.36C16:0 13.95 15.38 15.29 17.73 14.96 16.95C16:1n-7 3.08 2.83 3.48 3.92 3.13 3.72C17:0 0.21 0.19 0.24 0.16 0.12 0.14C18:0 4.56 4.42 4.85 4.45 4.83 5.40C18:1n-9 17.24 17.87 14.19 10.55 17.71 17.63C18:1n-7 3.01 2.80 3.17 3.31 3.21 3.40C18:2n-6 27.99 29.69 26.62 24.18 25.62 20.77C18:3n-3 4.29 3.86 3.98 3.26 3.56 2.80C20:1n-9 5.05 4.94 5.70 6.74 5.31 5.95C20:4n-6 0.61 0.40 0.60 0.63 0.79 1.01C20:5n-3 5.73 5.58 6.06 7.43 6.49 6.80C22:1n-9 3.24 2.96 4.00 4.62 3.30 3.53C22:6n-3 8.12 7.37 8.20 9.40 8.50 8.70 ∑ SFA a 20.54 21.89 22.45 24.78 21.87 24.85 ∑ MUFA b 25.30 25.61 23.06 20.6 26.23 26.98 ∑ PUFA c 32.28 33.55 30.60 27.44 29.18 23.57 ∑ HUFA d 14.46 13.35 14.86 17.46 15.78 16.51 ∑ n-3 e / ∑ n-6 f  0.63 0.56 0.67 0.81 0.70 0.84 a ∑ SFA, saturated fatty acid: C16:0, C14:0, C18:0, C17:0. b ∑ MUFA, monounsaturated fatty acids: C16:1n-7, C18:1n-9, C18:1n-7, C20:1n-9. c ∑ PUFA, polyunsaturated fatty acid: C18:2n-6, C18:3n-3. d ∑ HUFA, highly unsaturated fatty acids: C20:4n-6, C20:5n-3, C22:5n-3, C22:6n-3. e ∑ n-3:18:3n-3, 20:5n-3, C22:5n-3, 22:6n-3. f  ∑ n-6:18:2n-6, 20:4n-6.168  X. Li et al. / Aquaculture 430 (2014) 166  – 172  Moreover, ∑ HUFA and C22:6n-3 (DHA) levels increased accompany-ing the increasing EL supplementation. ∑ HUFA ranged from 28.72 ±3.40 to 33.77 ± 1.22, and was signi 󿬁 cantly higher in PL5 than inPL2 ( P   b  0.05). The n-3/n-6 ratio increased from 1.54 ± 0.10 (PL0) to2.02 ± 0.15 (PL5) with PL supplementation, and signi 󿬁 cantly higherin PL5 than in other groups ( P   b  0.05).The fatty acid composition (percent total fatty acids) of hepatopan-creaswaspresentedinTable7.Thefattyacidcompositionofhepatopan-creaswassimilartothoseinmuscles.The ∑ HUFAcontentrangedfrom9.43 ± 1.66 (PL1) to 14.44 ± 0.70 (PL5) ( P   b  0.05), while ∑ PUFA de-creased from 25.77 ± 2.38 (PL1) to 18.44 ± 0.46 (PL5) ( P   b  0.05).Crabs fed diet with 2% EL supplementation had the highest C20:4n-6,C22:6n-3 and C20:5n-3 percentage. The ∑ HUFA level was highest ingroup PL5, and signi 󿬁 cantly higher than in the PL0, PL1, PL3 and PL4groups ( P   b  0.05). 4. Discussion It has been demonstrated that dietary PL is utilized as a source of nutrient for early stages of   󿬁 sh and crustacean (Coutteau et al., 1997).Several papers have also reported that diets supplemented with PL could enhance the ef  󿬁 ciency of lipid utilization and supply preformedPC for growth (Sánchez et al., 2012; Tocher et al., 2008). The studies of  juvenile  L. vannamei  (Gong et al., 2000; González-Félix et al., 2002b)and juvenile  Penaeus monodon  (Kumaraguru vasagam et al., 2005)showed that growth was signi 󿬁 cantly increased with dietary PL   Table 3 Growth performances, feed conversion ratio, hepatosomatic index of juvenile  Portunus trituberculatus  fed diets containing different phospholipids sources and levels.Experiment diets (phospholipids %)Treatment PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%)Survival 76 ± 7 72 ± 11 66 ± 17 70 ± 7 67 ± 4 67 ± 6IBW 1 22.23 ± 0.05 22.38 ± 0.10 22.23 ± 0.08 22.19 ± 0.16 22.19 ± 0.22 22.14 ± 0.12FBW 2 69.66 ± 1.06 a 78.48 ± 3.96 bc 73.47 ± 1.00 ab 76.47 ± 1.14 b 78.92 ± 2.87 bc 83.11 ± 5.22 c WG 3 213.43 ± 5.48 a 250.65 ± 18.14 bc 230.45 ± 5.43 ab 244.57 ± 2.68 b 255.65 ± 13.86 bc 275.28 ± 21.71 c SGR  4 2.00 ± 0.03 a 2.20 ± 0.09 b 2.10 ± 0.03 ab 2.17 ± 0.01 b 2.23 ± 0.07 bc 2.32 ± 0.10 c FCR  5 2.04 ± 0.64 2.07 ± 0.6 2.00 ± 0.39 1.94 ± 0.32 1.54 ± 0.10 1.46 ± 0.09DFI 6 0.72 ± 0.04 b 0.68 ± 0.03 ab 0.70 ± 0.05 b 0.71 ± 0.01 b 0.68 ± 0.02 ab 0.65 ± 0.05 a HSI 7 5.83 ± 0.25 6.02 ± 0.38 5.28 ± 0.71 5.85 ± 1.12 5.83 ± 0.74 5.09 ± 0.71MF 8 1.43 ± 0.30 1.70 ± 0.15 1.57 ± 0.20 1.63 ± 0.44 1.79 ± 0.16 1.67 ± 0.14Data are represented as mean ± SD ( n  = 3). Values in a same column that do not share same superscripts are signi 󿬁 cantly different ( P   b  0.05). 1 Initial body wet weight (g). 2 Final body wet weight (g). 3 Weight gain = 100 × ( 󿬁 nal body weight − initial body weight) / initial body weight (g). 4 Special growth ratio = 100 × (LN  󿬁 nal weight − LN initial weight) / 56 days. 5 Feed conversion ratio = g total feed / g wet weight gain. 6 Daily feed intake = 100 × total feed / ( 󿬁 nal body weight + initial body weight) × 2 × 56 days. 7 Hepatosomatic index = 100 × hepatopancreas wet weight (g) /  󿬁 sh wet weight (g). 8 Molting frequency = 2 × molting / (initial crabs +  󿬁 nal crabs).  Table 4 Effects of experimental diets on serum cholesterol, glucose, triglycerides, and SOD levels in juvenile  Portunus trituberculatus. Treatment Cholesterol (mg/dl) Glucose (mmol/l) Triglycerides (mg/dl) SOD (U/ml)PL0 (0%) 3.98 ± 1.42 ab 3.69 ± 0.22 0.73 ± 0.05 101.61 ± 8.34PL1 (SL, 1%) 5.87 ± 1.71 b 3.64 ± 0.80 0.85 ± 0.11 95.78 ± 16.10PL2 (SL, 2%) 4.08 ± 0.62 ab 4.09 ± 0.86 0.99 ± 0.10 86.05 ± 15.93PL3 (SL, 4%) 3.57 ± 1.18 a 3.06 ± 0.17 0.77 ± 0.11 100.73 ± 13.02PL4 (EL, 1%) 4.78 ± 0.38 ab 2.98 ± 1.12 0.88 ± 0.25 100.20 ± 12.66PL5 (EL, 2%) 6.28 ± 1.39 b 3.91 ± 0.33 0.81 ± 0.19 90.65 ± 12.79Data are represented as mean ± SD ( n  = 3). Values in a same column that do not share same superscripts are signi 󿬁 cantly different ( P   b  0 .05).  Table 5 Whole body, muscle and hepatopancreas composition of juvenile  Portunus trituberculatus  fed diets containing different phospholipids sources and levels.Experiment diets (phospholipids %)Treatment PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%) Whole body Ash 8.23 ± 0.17 7.87 ± 1.12 7.47 ± 0.77 7.44 ± 0.12 7.51 ± 0.54 7.41 ± 0.54Moisture 73.06 ± 0.15 74.33 ± 0.02 74.05 ± 0.88 72.72 ± 2.60 74.90 ± 2.55 76.09 ± 4.62Protein 12.67 ± 1.29 10.85 ± 0.62 11.60 ± 0.72 11.63 ± 1.25 11.06 ± 1.39 10.58 ± 1.87Lipid 2.18 ± 0.46 1.88 ± 0.44 1.92 ± 0.26 2.16 ± 0.67 1.83 ± 0.55 1.89 ± 0.75 Breast muscle Moisture 80.66 ± 0.62 80.83 ± 2.28 75.92 ± 6.89 81.27 ± 0.79 79.62 ± 0.79 78.88 ± 1.38Protein 14.35 ± 0.72 14.80 ± 1.96 18.35 ± 5.90 13.78 ± 0.61 15.50 ± 0.53 16.34 ± 1.27Lipid 1.17 ± 0.20 ab 1.25 ± 0.22 ab 1.46 ± 0.19  b 1.00 ± 0.15 a 1.05 ± 0.16 ab 1.28 ± 0.32 ab Hepatopancreas Moisture 68.13 ± 2.29 69.39 ± 6.39 67.44 ± 2.26 68.92 ± 3.32 64.59 ± 1.31 63.41 ± 3.52Protein 10.75 ± 1.03 9.84 ± 1.62 12.19 ± 0.77 10.80 ± 2.45 12.09 ± 3.75 13.85 ± 0.83Lipid 13.49 ± 0.65 13.67 ± 0.44 13.29 ± 1.21 14.20 ± 2.45 15.45 ± 1.73 15.35 ± 1.28Data are represented as mean ± SD ( n  = 3). Values in a same column that do not share same superscripts are signi 󿬁 cantly different ( P   b  0.05).169  X. Li et al. / Aquaculture 430 (2014) 166  – 172  supplementation.Similarity,resultsfrompresentstudyalsoshowedthatWG signi 󿬁 cantly increased with 1% SL supplementation ( P   b  0.05), butdecreased slightly at higher SL supplementation. The negative effects of excess dietary PL supplementation were also reported in previousstudies (Coutteau et al., 1996; Teshima et al., 1986b). Besides, crabs fedPL2 diet had relativelylower molting frequency and poorer growthper-formancethanthoseinPL1andPL3groups,butnosigni 󿬁 cantdifferenceswere observed among these groups, and the reason is still not clearunder this experiment condition. Moreover, WG was higher in crabsfed diets with 1% or 2% EL than those fed control diet ( P   b  0.05), but nosigni 󿬁 cant differences was observed among crabs fed EL supplementa-tion diets. Based on growth performance, this study suggested thatdiets supplemented with 1% PL (SL or EL) could meet the requirementof the juvenile  P. trituberculatus . This result was in agreement withother feeding investigations of crustaceans, such as 1 – 2% (purity 60%)for  P. merguiensis  (Thongrod and Boonyaratpalin, 1998), and 1.25% for Penaeus penicillatus  and  P. monodon  (Chen, 1993; Chen and Jenn, 1991).MarinecrustaceansneedmoreHUFA(C20:5n-3andC22:6n-3)thanPUFA (C18:3n-3 and C18:2n-6) (Kanazawa, 1993; Lim et al., 1997; Xuet al., 1994). It also has been demonstrated that the EL contains higher  Table 6 Selected fatty acid composition of juvenile  Portunus trituberculatus  breast muscle fed diets containing different phospholipids sources and levels (% of total fatty acids).Experiment diets (phospholipids %)Fatty acid PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%)C14:0 0.86 ± 0.14 0.93 ± 0.03 0.98 ± 0.14 1.07 ± 0.17 0.91 ± 0.15 0.97 ± 0.13C16:0 15.84 ± 0.66 a 16.83 ± 0.22 ab 16.00 ± 0.86 a 18.32 ± 0.36 c 16.79 ± 0.24 ab 17.41 ± 0.60 bc C16:1-7 1.03 ± 0.27  a 1.07 ± 0.05 a 1.28 ± 0.28 ab 1.72 ± 0.38 b 1.09 ± 0.32 a 1.20 ± 0.07 a C17:0 0.47 ± 0.09 0.55 ± 0.19 0.44 ± 0.21 0.70 ± 0.16 0.51 ± 0.11 0.66 ± 0.26C18:0 8.74 ± 0.39 ab 8.69 ± 0.63 ab 7.40 ± 1.41 a 9.02 ± 0.22 b 9.33 ± 0.70 b 9.15 ± 0.30 b C18:1n-9 20.19 ± 0.54 19.67 ± 0.34 21.90 ± 3.57 19.25 ± 1.00 18.84 ± 1.88 19.85 ± 0.41C18:2n-6 18.02 ± 0.49 d 17.53 ± 0.83 cd 17.09 ± 0.79 bcd 16.40 ± 0.61 bc 15.91 ± 0.73 b 14.22 ± 0.79aC18:3n-3 1.04 ± 0.13 ab 1.01 ± 0.07 b 1.61 ± 0.61ab 1.09 ± 0.14ab 0.96 ± 0.15 ab 0.94 ± 0.07aC20:1n-9 1.18 ± 0.75 a 0.91 ± 0.16 ab 2.06 ± 1.06 b 1.40 ± 0.23 b 1.11 ± 0.57 ab 1.46 ± 0.47bC20:2n-7 1.15 ± 0.04 1.13 ± 0.21 1.03 ± 0.38 1.05 ± 0.03 1.15 ± 0.10 1.02 ± 0.14C20:4n-6 1.59 ± 0.43 ab 1.37 ± 0.22 a 1.26 ± 0.02 a 1.50 ± 0.02 a 1.73 ± 0.14 ab 1.99 ± 0.13 b C20:5n-3 15.70 ± 1.35 15.27 ± 1.44 15.19 ± 1.58 15.59 ± 1.32 16.68 ± 1.38 17.47 ± 0.62C22:6n-3 13.40 ± 0.52 ab 13.35 ± 0.70 ab 12.27 ± 2.08 a 13.42 ± 0.92 ab 13.86 ± 0.34 ab 14.30 ± 0.49 b ∑ SFA 1 25.92 ± 0.33 ab 27.00 ± 0.77 bc 24.82 ± 2.44 a 29.11 ± 0.61 c 27.53 ± 0.38 bc 28.19 ± 0.66 c ∑ MUFA 2 22.87 ± 1.46 22.21 ± 0.49 25.68 ± 4.56 23.07 ± 1.16 21.55 ± 2.37 23.27 ± 0.62 ∑ PUFA 3 19.06 ± 0.62 d 18.55 ± 0.89 cd 18.70 ± 0.94 cd 17.49 ± 0.75 bc 16.87 ± 0.71 b 15.16 ± 0.75 a ∑ HUFA 4 30.70 ± 2.17 ab 29.99 ± 2.31 ab 28.72 ± 3.40 a 30.51 ± 2.22 ab 32.26 ± 1.75 ab 33.77 ± 1.22 b ∑ n-3 5 / ∑ n-6 6 1.54 ± 0.10 a 1.57 ± 0.13 ab 1.58 ± 0.15 ab 1.68 ± 0.08 ab 1.79 ± 0.15 b 2.02 ± 0.15 c Data are represented as mean ± SD ( n  = 3). Values in a same column that do not share same superscripts are signi 󿬁 cantly different ( P   b  0.05). 1 ∑ SFA, saturated fatty acid: C16:0, C14:0, C18:0, C17:0. 2 ∑ MUFA, monounsaturated fatty acids: C16:1n-7, C18:1n-9, C18:1n-7, C20:1n-9. 3 ∑ PUFA, polyunsaturated fatty acid: C18:2n-6, C18:3n-3. 4 ∑ HUFA, highly unsaturated fatty acids: C20:4n-6, C20:5n-3, C22:5n-3, C22:6n-3. 5 ∑ n-3:18:3n-3, 20:5n-3, C22:5n-3, 22:6n-3. 6 ∑ n-6:18:2n-6, 20:4n-6.  Table 7 Selected fatty acid composition of juvenile  Portunus trituberculatus  hepatopancreas fed diets containing different phospholipids sources and levels (% of total fatty acids).Experiment diets (phospholipids %)Fatty acid PL0 (0%) PL1 (SL, 1%) PL2 (SL, 2%) PL3 (SL, 4%) PL4 (EL, 1%) PL5 (EL, 2%)C14:0 1.64 ± 0.16 1.64 ± 0.20 1.76 ± 0.06 1.99 ± 0.29 1.60 ± 0.23 1.78 ± 0.38C15:0 0.40 ± 0.14 0.37 ± 0.22 0.46 ± 0.02 0.47 ± 0.05 0.33 ± 0.06 0.39 ± 0.05C16:0 16.74 ± 1.64 a 18.54 ± 1.65 a 16.47 ± 0.34 ab 19.70 ± 0.50 b 17.35 ± 1.11 a 18.46 ± 1.31 ab C16:1n-7 3.16 ± 0.55 ab 2.92 ± 0.01 ab 2.83 ± 0.54 a 4.51 ± 1.30 b 3.23 ± 0.66 ab 3.92 ± 1.25 ab C17:0 0.56 ± 0.08 0.39 ± 0.24 0.53 ± 0.04 0.54 ± 0.09 0.40 ± 0.13 0.50 ± 0.17C18:1n-9 29.89 ± 1.84 27.87 ± 0.70 28.37 ± 0.68 26.28 ± 2.27 28.79 ± 4.71 30.18 ± 0.19C18:2n-6 22.05 ± 0.56 c 22.01 ± 1.14 c 21.00 ± 0.39 bc 17.80 ± 1.34 a 19.87 ± 1.34 b 16.65 ± 0.50 a C18:3n-3 3.01 ± 0.23 ab 3.76 ± 2.35 b 2.49 ± 0.06 ab 2.17 ± 0.44 ab 2.17 ± 0.17 ab 1.80 ± 0.26 a C20:1n-9 5.19 ± 1.79 a 6.45 ± 0.34 ab 7.23 ± 0.30 b 7.59 ± 0.24 b 6.65 ± 0.71 ab 7.78 ± 0.64 b C20:2n-7 1.54 ± 0.06 1.31 ± 0.10 1.64 ± 0.19 1.48 ± 0.01 1.37 ± 0.43 1.52 ± 0.22C20:4n-6 0.98 ± 0.18 ab 0.79 ± 0.16 a 0.96 ± 0.04 ab 1.06 ± 0.08 b 1.10 ± 0.05 b 1.41 ± 0.02 c C20:5n-3 4.41 ± 0.16 ab 3.07 ± 1.89 a 4.53 ± 0.09 ab 4.90 ± 0.38 b 4.14 ± 0.10 ab 5.07 ± 0.07 b C22:1n-9 2.54 ± 1.75 1.72 ± 2.00 3.01 ± 1.97 3.08 ± 2.06 2.31 ± 2.00 2.45 ± 2.31C22:6n-3 6.03 ± 0.34 a 5.57 ± 0.27 a 7.72 ± 0.32 b 7.35 ± 0.40 b 6.30 ± 0.85 a 7.96 ± 0.66 b C24:0 0.20 ± 0.02 0.69 ± 0.41 0.27 ± 0.01 0.52 ± 0.41 0.19 ± 0.05 0.59 ± 0.62 ∑ SFA 1 19.34 ± 1.55 a 20.94 ± 2.05 ab 19.22 ± 0.25 a 22.71 ± 0.35 b 19.68 ± 1.14 a 21.13 ± 1.30 ab ∑ MUFA 2 40.77 ± 2.07 ab 38.96 ± 2.78 a 41.44 ± 1.46 ab 41.46 ± 1.77 ab 40.98 ± 3.18 ab 43.51 ± 1.44 b ∑ PUFA 3 25.07 ± 0.45 d 25.77 ± 2.38 d 23.49 ± 0.40 cd 19.97 ± 0.97 ab 22.04 ± 1.49 bc 18.44 ± 0.46 a ∑ HUFA 4 11.42 ± 0.03 b 9.43 ± 1.66 a 13.21 ± 0.36 c 13.31 ± 0.22 c 11.54 ± 0.86 b 14.44 ± 0.70 c ∑ n-3 5 / ∑ n-6 6 0.58 ± 0.03 ab 0.55 ± 0.07 a 0.67 ± 0.01 bc 0.77 ± 0.09 cd 0.60 ± 0.07 ab 0.82 ± 0.06 d Data are represented as mean ± SD (n = 3). Values in a same column that do not share same superscripts are signi 󿬁 cantly different ( P   b  0.05 ) 1 ∑ SFA, saturated fatty acid: C16:0, C14:0, C18:0, C17:0; 2 ∑ MUFA, monounsaturated fatty acids: C16:1n-7, C18:1n-9, C18:1n-7, C20:1n-9; 3 ∑ PUFA, polyunsaturated fatty acid: C18:2n-6, C18:3n-3; 4 ∑ HUFA, highly unsaturated fatty acids: C20:4n-6, C20:5n-3, C22:5n-3, C22:6n-3; 5 ∑ n-3:18:3n-3, 20:5n-3, C22:5n-3, 22:6n-3; 6 ∑ n-6:18:2n-6, 20:4n-6.170  X. Li et al. / Aquaculture 430 (2014) 166  – 172
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