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Lipid class distribution and fatty acid composition of wild and farmed prawn, Penaeus monodon (Fabricius)
65
Aquaculture, 89 (1990) 65-81 Elsevier Science Publishers B.V., Amsterdam
Lipid class distribution and fatty acid cornposition of wild and farmed prawn, Penaeus monodon ( Fabricius )
Cliona D. O’Leary” and Anthony D. Matthewsb aOverseasDevelopment Natural Resources Institute, Central Avenue, Chatham Maritime, Chatham, Kent, ME4 4TB (Great Britain) bBio,compatiblesLtd., Brunel Science Park, Kingston Lane, Uxbridge, Middlesex, UB8 3PQ (Great Britain) (Accepted 10 January 1990 )
ABSTRACT O’Leary, (CD. and Matthews, A.D., 1990.Lipid class distribution and fatty acid composition and farmed prawn, Penaeus monodon (Fabricius). Aquaculture, 89: 65-8 1.
of wild
Total lipid levels of the muscle of wild and farmed Penaeus monodon were similar (4.35% and 4.66% dry weight) whilst head lipid levels (hepatopancreas) were higher in farmed P. monodon (farmed !).36%, wild 7.07%). The lipids of the muscle of wild P. monodon are mainly composed of phospholipid (70.5%), triglyceride (2.9%) and cholesterol (19.84Oh). Head lipids have higher triglyceride levels (9.18%) and lower levels of phospholipid (56.7Oh). Farmed P. monodon muscle lipids had lower levels of phospholipid (57.0%) and higher levels of triglyceride (8.46%) and cholesterol (24.17%) in comparison to wild P. monodon. Head lipids also showed lower phospholipid (39.23%) and higher triglyceride (25.93%). The composition of the phospholipids of wild and farmed P. monodon was similar excepting spingomyelin. The major fatty acids of the muscle phospholipids of wild P. monodon were 16:O (13.0%), 16:l (5.3%), 18:O (13.2%), 18:ln-9 (8.4%) 18:2n-6 (1.9%), 20:4n-6 ( 12.9%), 20: 5n-3 (13.6%) and 22: 6n-3 (15.5%). Head phospholipids had almost the same profile. Farmed P. monodon phospholipid fatty acids were similar except that levels of 18 : 2n-6 ( 9.8%) were higher and 16: 1 ( 1.79%) and 20:4n-6 (4.9%) lower. The lipid composition of hatchery and wild post-larvae (approximately post-larvae day 20) was dissimilar. Wild post-larvae lipid was almost entirely phospholipid (68.7%) and cholesterol (27.8%) whilst hatchery stock showed lower phosphohpid levels and significant amounts of triglyceride (5.9-15.6%). Comparison of the fatty acids of the phospholipids showed that wild post-larvae had lower levels of 18: ln-9, 18: 2n-6 and higher levels of 16: 1 and 22:6n-3 (20.4% wild, 4.0% farmed). Analysis of diets fed to farmed P. monodon showed low levels of phospholipid and very high levels of free fatty acids, indicative of dietary lipid deterioration. Diets used for P. monodon are those which have been developed for P. japonicus. Differences, however, exist in the lipid composition and fatty acid profiles of the two wild stocks suggesting differing lipid formulations may be required in diets.
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66
CD.
O’LEARY
AND
A.D.
MATTHEWS
INTRODUCTION
Culture of the tiger prawn, Penaeus monodon is increasing substantially in subtropical and tropical countries. Whilst many papers have been presented concerning the nutritional requirements of Penaeus juponicus, those for Penaeus monodon are sparse and specific dietary requirements for this species have not been established. Phospholipids have been shown to be essential in the diets of penaeids (Kanazawa et al., 1985; Teshima et al., 1986) as have the highly unsaturated fatty acids 20 : 5n-3 and 22 : 6n-3 (Kanazawa et al., 1979a). Whilst phospholipids are almost certainly involved in lipid and cholesterol transport in the prawn (Teshima et al., 1986) they are also the main structural component of cell membranes and correct formation of these membranes is necessary in order that the cells may function effectively. Many intensive commercial farms in Indonesia have reported stressed or weak stock (personal communications to A.D. Matthews) and whilst numerous factors may be involved, the overwhelming importance of dietary lipids and particularly phospholipids and the absence of basic lipid information for P. monodon, led to the present study. MATERIALS AND METHODS
Samples of wild and farmed P. monodon were collected from fishermen and farms along the coastline near Jepara, Central Java, Indonesia, during November 1988. Farmed samples, large wild P. monodon and hatchery and wild post-larvae samples were obtained live and transported on ice to the marine laboratory of the University of Diponegoro, Jepara, for immediate analysis. Small wild P. monodon (less than 30 g ) were obtained un-iced from the lishing boats. Diets were purchased from local suppliers. Post-larvae were analysed whole. For all other samples, heads were broken from the tail and tail muscle was extracted from the carapace before analysis. Lipids were extracted by the method of Pozo et al. ( 1988 ) . Lipids were separated into class by thin layer chromatography on high performance silica gel plates, layer thickness 200 pm, having a preabsorbent layer (Whatman) using methyl acetate, propan- l-01, chloroform, methanol, KC1 (25 : 25 : 10 : 9 : 0.25%) as the developing solvent for phospholipids and hexane, diethyl ether, acetic acid ( 75 : 23 : 2 ) as the developing solvent for non-polar materials. The developed plates were sprayed with 8% orthophosphoric/cupric spray and charred in an oven at 180°C for 10 min. Quantification of lipid fractions was by densitometry ( Joyce-Loebl Chromoscan 3 ) . Phospholipid fractions for gas-liquid chromatography (GLC) were isolated by silicic acid column chromatography (Ramsey and Patterson, 1948 ). Fatty acid methyl esters were prepared by the method of Metcalfe and Wang ( 198 1) and separated by GLC on a Supel-
LIPIDS AND I=ATTY ACIDS IN PENAEUS MONODON
61
cowax 10 fused silica capillary column. Fatty acids were identified by comparison with a polyunsaturated fatty acid standard, marine source (Supelco, U.K.). RESULTS
Total lipid and lipid class distribution Total muscle lipid levels (Tables 1 and 2) of wild and farmed P. monodon were similar, ranging from 3.8 to 4.7% dry weight wild (average 4.35%) and 3.5 to 5.4% farmed (average 4.66%). Head total lipid levels were higher in farmed (average 9.36%) than wild (average 7.07%). The lipid class distribution (Tables 3 and 4) shows this extra lipid is triglyceride. Wild muscle had a higher level of phospholipid (wild average 70.54%, farmed 57.0%) and lower triglyceride (wild average 2.9%, farmed 8.46%) and a lower level of chlolesterol (wild average 19.84%, farmed 24.17%). The increase in cholesterol with decreasing phospholipid concentrations may be significant. Postlarvae (Table 5 ) total body lipids showed a similar pattern. Total lipid levels of the diets (Table 6 ) were within the range generally utilized for penaeids of 6-l 0%. Phospholipid levels in the diets were low (in general N 1.25%). This may be due to the breakdown of phospholipid since free fatty acid levels were very high (average 24.1% in the lipid). Cholesterol levels in the diets fall within th.e optimal range, 0.5-1.0% (Kanazawa, 1985). TABLE 1 Moisture and total lipid levels of wild P. monodon Sample no.” -
Sex
Head Muscle
1 3 5 7 9b 10”
2 4 6 8
11 12 13 14
Number of prawns in sample 16 14 5 3 2 2
male female female male
1 1 1 1
Whole body Weight (g)
lo.oei 1.7 16.9 X!I 3.7 37.7 + 10.2 81.1 f 5.5 98.1 139.7 152.1 169.3
Length (cm) 11.5t0.6 13.7kO.9 17.0f 1.1 21.8+ 1.0
24.8 27.5 27.5 28.0
Moisture (O/o)
Lipid (%)
Head Muscle
Head ~-
77.7 77.6 75.6 74.1 74.1 72.1
78.5 77.3 76.7 76.1
74.6 73.6 73.2 74.6
Muscle
wwd dw ww
dw
1.6 1.8 1.8 1.7 1.6 1.9
0.9 1.0 1.0 0.9
4.2 4.4 4.3 3.8
1.2 1.1 1.2 1.2
4.7 4.2 4.5 4.7
7.2 8.0 7.4 6.8 6.2 6.8
‘Samples l-4 and 5- 14 were obtained from the fishing villages of Wedung and Rembang, respectively. bHeads from samples 11 and 14. “Heads from samples 12 and 13. ?vw= wet weight, dw = dry weight. “Results as shown are mean k s.d.
16 18 20 22 24 26 28 30 32 34 36 38
15 17 19 21 23 25 27 29 31 33 35 37
15 44 20 50 1 lb 27 9 15 20 25
I
Stocking density ( nlm2)
3 6 6 6 6 8 not known 12 10 12 16 18
Age’ (weeks)
none A,B E C C none none E B A,C D C
Dietd
28 10 12 8 7 10 11 4 7 16 7 5
Number of prawns in sample
1.8+0.8 2.3+ 1.0 2.8k2.1 3.5 f 0.9 3.6t0.7 9.6f2.5 1 l.Ok3.0 12.6f4.7 14.0f2.3 19.0f4.9 39.2t7.3 42.0 f 9.9
6.5* I.1 7.1+1.0 7.321.8 8.2+ 1.2 8.5f0.5 11.6+1.0 12.2& 1.1 12.4+ 1.5 12.8kO.7 14.6+ 1.2 18.0+_ 1.1 18.6+ 1.3
74.5 76.6 74.3 72.1 74.4 72.6 75.7 75.9
78.3 77.0 74.2 75.8 76.0 73.7 75.9 77.4 2.6 0.8 2.5 1.6 1.5 1.9 2.7 3.2 3.0 2.4 2.0
9.8 6.4 7.4 9.7 12.5 10.9 9.9 8.3
1.2 1.4 1.2 1.3 0.8 1.0 1.3 1.3 1.2 1.1 1.0
ww
Ww
dw
Muscle
Head
Muscle
Head
Weight (g)
Length (cm)
Lipid (%)
Moisture (%)
Whole body
5.5 3.5 3.9 5.4 5.4 4.6 4.6 4.4
dw
“Samples 15-18, 25, 26, 33-36 were obtained from The Brackishwater Aquaculture Development Centre, Jepara: samples 21, 22, 27, 28, 37 and 38 from private farms near the village of Bulak Baru; samples 19, 20, 23,24 and 30 from a private farm near the village of Bondo and samples 3 l-32 from a private farm, Semarang. bWild post-larvae, all others hatchery stock. ‘Age beyond PL20. dSee Table 6.
Muscle
Head
Sample no.a
Moisture and total lipid levels of farmed P. monodon
TABLE 2
69
LIPIDS AND FATTY ACIDS IN PENAECJS MONODON
TABLE 3 Percentage lipid class distribution
of wild P. monodon
Sample no.
PL
TG
CHOL
FFA
MG/DG
CE
Muscle 2 4 6 8 11 12 13 14 Average
62.2 62.3 69.7 79.5 72.1 78.0 70.7 69.8 70.54+ 5.89
0.0 1.4 2.8 0.0 4.5 0.0 4.2 10.3 2.9 k 3.29
19.0 19.7 19.3 18.8 21.5 21.8 19.7 18.9 19.84f 1.10
8.3 10.9 6.7 0.0 0.0 0.0 0.0 0.0 3.24f4.31
0.0 3.1 1.5 1.7 1.9 0.2 5.4 1.0 1.852 162
10.5 2.6 0.0 0.0 0.0 0.0 0.0 0.0
48.2 49.2 60.5 54.6 62.3 65.4 56.7f6.51
8.0 14.6 9.3 6.0 10.0 7.2 9.18k2.75
22.6 14.2 17.3 22.1 19.0 18.8 19.Ok2.84
17.6 17.6 8.3 9.1 8.2 6.7 11.25f4.55
0.0 4.4 4.6 8.2 0.5 1.9 3.27 k2.82
3.6 0.0 0.0 0.0 0.0 0.0
Head 1 3 5 7 9 10 Average
“PL=phospholipid; TG= triglyceride; CHOL=cholesterol; monoglyo:rides/diglycerides; CE = cholesterol esters.
FFA=free
fatty
acids;
MG/DG=
Composition of the phospholipids The composition of the phospolipid of wild and farmed P. monodon (Tables 7 and 8 ) in respect of phosphatidylcholine, phosphatidylserine, phosphatidylinositol and phosphatidylethanolamine were not found to be significantly different. Sphingomyelin levels were, however, higher in wild muscle and head phospholipids (wild muscle 15.0%, head 9.03%; farmed muscle 11.39%, head 7.36%). Fatty acid composition of the phospholipidsof P. monodon The major fatty acids of both wild and farmed muscle and head phospholipidwere foundto be 16:0, 16: 1, 18:0, 18: ln-9, 18:2n-6,20:4n-6,20:5n-3 and 22 : 6n-3 (Tables 9, 10 and 11). The only major differences were in levels of 16 : 1 (wild muscle average 5.3%, head 5.6%; farmed average 1.7%, 2.3%), 18.2n-6 (wild muscle 1.901’0, head 1.8%; farmed 9.8%, 9.6%) and 20 : 4n-6 (wild muscle 12.9%, head 12.8%; farmed 4.9%, 5.4%). The same fatty acids (Table 12) dominate the fatty acid profiles of wild and farmed post-larvae phospholipids but hatchery stock had higher levels of 18 : 2n-6, 18 : 3n-3 and a low level of 22: 6n-3. The fatty acid profiles for the range of diets made by any one manufacturer were found to be similar (Table 13 ). Differences between products in the
70
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 4 Percentage lipid class distribution
of farmed P. monodon
Sample no.
PL”
TG
CHOL
FFA
Muscle 16 18 20 22 24 26 28 30 32 34 36 38 Average
42.4 49.9 60.5 48.6 68.9 38.9 67.8 49.6 69.5 62.5 61.4 64.1 57.0f 10.18
20.8 24.0 0.0 6.7 0.0 22.3 2.7 6.9 0.0 11.6 6.5 0.0 8.46 f 8.78
22.0 16.2 36.8 20.9 31.0 20.0 23.7 24.6 30.5 17.3 17.3 29.8 24.17f6.25
10.6 7.0 0.0 14.0 0.0 9.7 0.0 11.1 0.0 5.1 5.4 0.0 5.24+ 5.00
4.2 2.9 2.7 9.8 0.1 9.1 5.8 7.8 0.0 3.5 9.4 6.1 5.1223.30
Head 15 17 19 21 23 25 27 29 31 33 35 37 Average
31.8 36.5 42.8 30.3 46.0 34.7 48.8 35.0 36.6 38.0 44.1 46.2 39.23 + 5.86
39.1 40.9 8.0 11.5 11.0 34.5 33.6 17.8 43.3 36.6 28.9 5.9 25.93 f 13.46
16.6 13.4 25.3 15.1 23.0 13.5 16.7 14.3 18.8 13.4 17.1 20.1 17.27k3.72
12.5 9.1 20.7 30.9 9.8 12.2 0.9 17.8 0.0 8.6 5.1 17.1 12.06 f 8.33
0.0 0.1 3.2 10.2 10.2 5.1 0.0 15.1 I.3 3.4 4.8 10.7 5.34t4.84
“For explanation of abbreviations
MG/DG
see Table 3.
TABLE 5 Lipid class distribution
of post-larvae of P. monodon
Sample no.
Origin
Diet
No. of prawns in sample
Postlarval day
PL”
TG
39
bPT Yayasan Dian Desa
Skeletonema, Artemia, Frippak
2000
20
54.6
10.3 35.1
40
bPT Alam Windu Tama
Skeletonemu, Taiwanese inert
1500
22
52.5
15.6 22.7
41
bPT Urang Baruna
Skeletonema, Taiwanese inert
1700
16
63.7
5.9 20.3
10.1
42
Benteng Portugis
Wild
2000
30”
68.7
0.4 27.8
3.1
“For explanation of abbreviations bHatchery. ‘Estimated.
see Table 3.
CHOL
MG/DG
0.0 9.2
71
LIPIDS AND FATTY ACIDS IN PENAEUS MONODON
TABLE 6 Moisture, total lipid and lipid class distribution Diet Name
Origin
Designation
A
Fanni
Taiwan
2 5
7.8 9.0
9.9 9.8
not done 12.5 40.6 11.3
20.4
9.5
5.7
B
Bama
Indonesia
0 1 2 3 4 5
7.1 9.9 9.3 9.8 12.0 10.2
8.8 9.4 7.2 7.2 7.2 5.2
17.5 17.8 14.4 14.1 14.2 18.7
39.1 7.6 38.4 9.6 35.3 9.6 36.7 10.1 32.0 9.4 31.5 9.3
18.8 18.1 21.0 22.6 22.0 21.6
11.9 11.8 15.1 12.7 19.2 15.9
5.1 4.3 4.6 3.8 3.2 3.0
C
Tungp ao
Taiwan
1A 2 3 4 5
10.8 10.6 10.9 11.5 10.4
9.5 8.1 8.1 7.0 7.5
11.8 11.3 15.2 15.8 16.9
34.9 9.8 41.9 6.3 36.5 13.3 28.1 16.1 30.0 15.5
28.0 12.7 25.4 9.3 22.3 11.4 27.4 10.5 23.1 11.3
2.8 5.8 1.3 2.1 3.2
D
Rajaudang
Indonesia
4 5
10.1 8.2
6.8 5.1
11.2 7.6
28.0 13.4 26.3 9.7
32.5 12.6 30.6 22.1
2.3 3.7
E
Chuen Shin Taiwan
1 2 3 3mths
10.7 11.0 8.4 11.0
7.4 8.0 6.7 6.0
18.5 42.6 7.3 15.1 47.5 8.3 not done 21.7 36.5 11.3
16.0 12.5 14.5 11.8 16.2 11.7
3.1 2.8
IA 1B 2 3 4 5
8.8 9.5 9.5 10.2 11.1 9.4
9.3 10.1 8.5 7.8 6.9 9.1
12.5 15.5 17.7 7.9 13.8 11.7
22.3 7.5 15.1 5.0 16.2 7.0 53.0 1.1 27.2 12.2 15.6 14.9
1.6 2.0 1.4 5.9 0.0 4.1
F
President Wanyng
Taiwan
aDry weighi. bFor explanation of abbreviations
Moisture (o/o)
of diets for post-larvae Total lipid” (Oh)
PLb (%)
TG
40.4 48.2 45.5 17.1 31.8 45.7
CHOL FFA MG/DG
15.7 14.2 12.2 15.0 15.0 8.0
CE
2.6
see Table 3.
range seem to be in pellet size only. Levels of 20 : 512-3and 22 : 6n-3 are at or above the level of 1% recommended by Kanazawa et al. ( 1979a). High levels of 18 : 2n-6 were found (presumably from soya lecithin) and low levels of 20 : 4~6. DISCUSSION
Total 1:ipidlevels of wild P. monodon muscle, average 1.1% wet weight (ww ), were similar to those reported for other penaeids (Guary et al., 1974; P. japnicus 1.2%; Gopakumar and Nair, 1975, P. indicus 1.3%; Chanmugan et al., 1983, P. aztecus 1.O%). Head lipid levels of wild P. monodon were also low,
72
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 7 Composition Sample no.
Muscle 2 4 6 8 11 12 13 14 Average
of the phospholipids Phospholipid”
of wild P. monodon
(%)
LysoPC
PC
PS
PI
3.0 1.5 0.0 0.9 0.0 0.0 0.0 0.0
43.0 44.8 43.3 44.0 47.1 44.4 46.2 45.9 44.84f
9.7 8.8 10.4 9.9 8.1 8.6 8.6 9.2 9.16f0.73
3.3 3.2 4.8 5.2 2.9 3.7 3.9 4.2 3.9kO.75
26.0 27.1 25.0 24.0 28.1 27.4 26.3 27.5 26.43 + 1.30
15.0 14.6 16.5 16.0 13.8 15.9 15.0 13.2 15.0f 1.05
2.8 2.2 2.8 3.8 2.8 2.9 2.88 + 0.47
27.0 25.4 27.5 26.4 27.6 28.4 27.05 20.9
8.4 11.0 9.8 9.4 7.8 7.8 9.03rt 1.16
1.36
PE
sphing
Head
1 3 5 7 9 10 Average
2.7 2.8 0.0 1.5 0.0 0.0
50.4 50.8 52.8 51.0 53.4 53.6 52.0& 1.3
a.7 7.8 7.1 7.9 8.4 7.3 7.87 + 0.56
“LysoPC = lysophosphatidylcholine; PC=phosphatidylcholine; PS=phosphatidylserine; phosphatidylinositol; PE=phosphatidylethanolamine; sphing=spingomyelin.
PI =
1.7% ww. Total body lipid levels of P. japonicus have been reported to be 2.8% ww with the hepatopancreas containing 10.5% lipid (Guary et al., 1974). It would appear, therefore, that hepatopancreas total lipid levels are considerably lower in P. monodon than in P. japonicus. Farmed and wild P. monodon had similar total muscle lipid contents but the lipid level of farmed P. monodon heads was higher, the extra lipid being triglyceride. Dietary imbalance in terms of the phospholipids : triglyceride ratio in the total lipid of the diets or carbohydrate: lipid ratios may be responsible. The lipid of wild P. monodon muscle and post-larvae was almost entirely phospholipid whilst both the muscle of farmed P. monodon and post-larvae from hatcheries showed reduced phospholipid and increased triglyceride. Wild P. monodon head lipid contained a significant amount of triglyceride but again phospholipid levels were lower in farmed P. monodon and triglyceride higher. Actual phospholipid concentrations were not signficantly different in the P. monodon heads (4.0% wild, 3.7% farmed) but were in the muscle. InsufIicient dietary phospholipid may have been the cause. The reported muscle lipid composition for P. japonicus (Guary et al., 1974) of 22.1% phospho-
LIPIDS AND F’ATTY ACIDS IN PENAEUSMONODON
73
TABLE 8 Composition Sample no.
Muscle 16 18 20 22 24 26 28 30 32 34 36 38 Average Head 15 17 19 21 23 25 27 29 31 33 35 37 Average
of the phospholipids Phospholipid”
of farmed P. monodon
(%)
LysoPC
PC
PS
PI
PE
sphing
0.0
45.1 48.0 50.4 52.6 49.7 40.8 47.2 46.2 51.0 50.8 48.0 42.7 47.71 f 3.39
8.3 9.0 1.2 1.9 6.7 12.1 8.1 10.1 4.1 7.3 8.5 8.9 8.23k 1.74
3.9 3.3 2.8 2.3 2.2 3.8 2.7 4.0 1.1 2.9 3.6 3.4 3.OkO.81
30.8 28.6 31.2 28.9 33.7 28.0 28.6 25.8 34.8 26.9 28.0 29.4 29.56 f 2.5
11.9 11.1 8.4 7.0 7.7 15.3 13.4 13.9 8.4 12.1 11.9 15.6 11.39k2.82
50.4 53.5 55.0 55.0 48.9 44.9 51.0 53.5 50.2 52.5 63.3 52.6 52.51 k4.22
6.6 7.2 6.9 5.2 8.9 11.4 9.2 8.5 8.6 9.3 8.2 7.9 8.16& 1.51
3.9 3.5 3.0 2.3 3.4 3.1 3.4 3.3 3.9 3.9 3.5 2.3 3.34f0.53
32.1 28.6 28.0 32.0 29.4 28.6 27.0 26.0 28.9 26.9 24.6 28.9 28.42 f 2.1
7.0 1.2 7.1 5.5 9.4 11.4 9.4 8.1 8.4 1.4 0.4 7.0 1.36 k 2.56
0.0 0.0
1.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
0.0 0.0 0.0 0.0 0.0 0.0
0.0 0.6 0.0 0.0 0.0 1.3
aFor explanation of abbreviations
see Table 7.
lipid, 4 1.5% sterols (cholesterol) and 12.2% triglyceride with a similar composition for ovary and hepatopancreas tissues is very different from the present results for P. monodon and may reflect differing dietary needs. The absence of de novo synthesis of linoleic ( 18 : 2n-6)) linolenic ( 18 : 3n3 ) , eicosapentaenoic (20 : 5n-3 ) and docosahexaenoic (22 : 6n-3 ). acids from acetate-‘“C or palmitic acid-14C has been demonstrated in P. japonicus (Kanazawa and Teshima, 1977; Kanazawa et al., 1979~) and P. monodon (Kanazawa et al., 1979d). Conversion of dietary linoleic acid to arachidonic acid and linolenic acid to eicosapentaenoic and docosahexaenoic acid has been observed but the conversion rates were slow (Kayama et al., 1980). Dietary
74
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 9 Fatty acid composition
of the phospholipids
Fatty acid Fatty acid composition
in samples of wild P. monodon
(O/o)
Muscle
Head
2
4
6
8
11
12
13
14
Av
1
3
5
7
9
10
Av
14:o 15:o 16:0 16:l 17:o 17:l 18:0 18: 1 n-9 18: 1 n-7 l8:2 n-6 18:3 n-3 20: 1 n-9 20: 1 n-7 20:2 20:4n-6 20:5 n-3 22:l n-11 22: 5 n-3 22:6n-3 24:1 Othersa
1.0 1.3 15.4 4.4 3.0 1.4 13.9 6.3 1.3 2.1 0.7 0.0 0.0 0.7 11.2 16.7 0.0 1.7 14.8 0.0 4.1
0.9 1.3 14.2 4.0 3.2 1.5 13.8 6.4 1.0 2.0 0.6 0.9 0.4 0.8 11.9 15.9 0.0 1.8 14.5 0.4 4.5
0.6 0.7 11.4 4.1 2.7 1.1 14.8 6.9 0.8 2.1 0.6 0.7 0.0 0.8 14.7 14.0 0.7 1.6 16.2 0.4 5.1
1.4 1.0 16.7 8.8 2.1 1.8 10.9 11.6 0.6 2.0 0.7 1.0 0.7 0.0 11.0 11.9 0.6 1.2 11.9 0.6 3.5
0.5 0.9 11.8 4.7 3.2 1.8 12.9 7.5 0.8 1.7 0.6 0.9 0.0 1.0 13.4 12.6 0.6 1.8 16.8 0.7 5.8
0.0 0.6 8.8 3.7 2.6 1.8 14.8 10.1 0.5 2.0 0.5 0.5 0.0 0.8 15.3 11.2 0.4 1.2 18.7 0.6 5.9
0.6 0.7 12.6 6.4 2.8 1.9 12.3 9.1 0.6 1.8 0.5 0.8 0.4 0.7 13.3 12.5 0.6 1.2 15.4 0.5 5.3
0.5 0.8 12.8 5.9 2.9 2.3 11.9 9.4 0.4 1.7 0.5 0.8 0.4 0.7 12.7 13.7 0.4 1.1 15.4 0.6 5.1
0.7 0.9 13.0 5.3 2.8 1.7 13.2 8.4 0.8 1.9 0.6 0.7 0.2 0.7 12.9 13.6 0.4 1.5 15.5 0.5 4.7
1.3 1.4 17.6 4.8 3.0 1.8 13.8 8.6 1.1 2.0 0.0 1.0 0.0 0.9 11.6 14.4 0.0 1.2 12.6 0.0 2.9
1.1 1.4 16.3 4.4 3.2 1.9 14.3 8.5 0.9 2.1 0.6 0.0 0.0 1.0 12.2 13.5 0.0 1.3 12.0 0.5 4.8
0.7 1.0 15.1 4.9 2.6 1.4 12.6 9.3 0.7 1.9 0.5 1.2 0.0 1.0 13.9 12.5 0.7 1.2 14.1 0.0 4.7
1.0 0.8 15.7 6.9 1.9 1.6 11.5 11.7 0.6 1.6 0.4 1.3 0.0 0.8 12.7 12.7 0.7 1.0 12.5 0.0 4.6
0.8 1.2 16.3 6.3 2.5 2.3 10.4 11.3 0.4 1.6 0.4 0.4 1.2 1.1 12.4 11.3 0.4 0.9 12.7 0.0 6.1
0.7 0.9 14.8 6.3 2.4 2.0 11.6 11.4 0.0 1.8 0.4 0.3 1.2 1.1 14.1 10.8 0.5 1.5 12.6 0.0 5.6
0.9 1.1 16.0 5.6 2.6 1.8 12.4 10.1 0.6 1.8 0.4 0.7 0.4 1.0 12.8 12.5 0.4 1.2 12.8 0.1 4.8
n-6PUFA n-3PUFA n-3/n-6
13.3 13.9 16.8 13.0 15.1 17.3 15.1 14.4 14.8 13.6 14.3 15.8 14.3 14.0 15.9 14.6 33.9 32.8 32.4 25.7 31.8 31.6 29.6 30.7 31.2 28.2 27.4 28.3 26.6 25.3 25.3 26.9 2.5 2.4 1.9 2.0 2.1 1.8 2.0 2.1 2.1 2.1 1.9 1.8 1.9 1.8 1.6 1.8
“Not identified.
sources of 20: k-3 and 22: 6n-3 are considered essential (Kanazawa, 1984) but not 20 : 4~6. In Table 14 the levels of the major fatty acids of P. monodon wild, P. japonicus, dietary components and commercial diets are compared. The major differences found between wild P. monodon and P. japonicus were in the fatty acids 20: 4n-6 and 20: ln-9. This may be a reflection of differing synthesis abilities, diet or environment. The fatty acid composition of the phospholipid of wild and farmed P. monodon was similar except for lower levels of 16: 1 and 20: 4n-6 and higher levels of 18 : 2n-6 in farmed. Adequate amounts of the essential fatty acids 20 : k-3 and 22 : 6n-3 seem to be provided by the diets. Phospholipids are the major component of biomembranes and are characterised by having a high content of polyunsaturated fatty acids that confer fluidity to the membranes. It has been demonstrated that fatty acid compo-
75
LIPIDS AND FATTY ACIDS IN PENAEUSMONODON
TABLE 10 Fatty acid composition Fatty acid
of the phospholipids
Fatty acid composition
in samples of farmed P. monodon muscle
(O/o)
16
18
20
22
24
26
28
30
32
34
36
38
Av
14:o 15:o 16:0 16:l 17:o 17:l 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9 20: 1 n-7 2012 20:4 n-6 20: 5 n-3 22:l n-11 2215 n-3 2216 n-3 24:l Others=
0.0 0.6 15.5 3.1 2.4 0.9 11.7 3.7 4.5 0.0 7.8 1.1 0.0 0.7 0.0 0.5 6.5 26.4 0.0 1.2 10.7 0.8 1.9
0.0 0.9 16.4 1.2 2.7 0.9 7.9 7.7 1.4 0.0 9.0 0.0 0.0 1.1 0.0 0.0 5.9 17.7 0.0 1.1 21.0 4.0 1.1
0.0 0.6 23.5 1.8 1.5 6.0 7.5 10.1 1.5 0.0 13.9 0.6 0.0 1.7 0.0 1.1 2.7 12.2 0.7 1.3 13.5 3.8 1.4
0.0 0.9 17.2 1.6 2.3 0.9 7.9 8.9 1.3 0.0 9.4 0.0 0.0 0.9 0.0 0.9 4.5 15.6 0.5 1.6 17.9 4.1 3.6
0.0 0.7 19.7 1.5 1.3 1.1 7.1 9.6 1.8 0.0 9.4 0.8 0.0 1.8 0.0 1.3 3.4 13.8 0.0 1.1 17.6 5.5 2.3
0.8 2.2 17.9 3.4 3.8 2.9 10.8 7.3 1.5 0.0 5.7 1.0 0.0 1.5 0.0 0.0 8.6 18.7 0.0 1.0 11.4 1.2 0.3
0.6 0.7 17.1 2.4 2.2 1.1 11.0 7.8 2.2 0.0 7.5 0.6 0.0 0.5 0.4 0.7 7.4 17.3 0.0 1.0 15.6 0.6 3.3
0.0 0.0 19.2 1.2 1.1 0.0 10.1 9.6 1.0 0.0 13.7 0.5 0.5 1.5 1.1 0.0 3.3 14.8 0.6 1.0 18.8 2.0 0.0
0.2 0.0 21.1 0.3 1.3 0.0 10.4 9.4 0.6 0.2 11.6 0.4 0.0 0.8 0.0 0.7 4.5 16.8 0.5 0.8 18.8 0.6 1.0
0.0 0.0 15.1 0.8 1.1 0.4 10.5 6.8 1.1 0.0 9.4 0.4 0.0 1.1 0.0 .08 3.8 18.5 0.3 1.0 27.3 0.6 1.0
0.0 0.2 17.7 1.6 1.0 0.4 9.2 8.9 0.8 0.0 8.8 0.3 0.0 1.7 0.0 0.7 3.9 16.9 0.4 0.8 23.0 1.9 1.6
0.0 0.0 19.7 1.5 1.1 0.6 11.6 13.6 0.8 0.0 10.9 0.0 0.0 0.7 0.0 0.9 4.4 16.2 0.0 0.8 16.0 1.1 0.0
0.1 0.5 18.3 1.7 1.8 0.8 9.6 8.6 1.5 9.8 0.5 1.2 0.1 0.6 4.9 17.1 0.3 1.1 16.1 2.2 -
n-6 PUFA n-3 PUFA n-3/n-6
14.3 39.4 2.8
14.9 39.8 2.7
16.6 27.6 1.7
13.9 35.1 2.5
12.8 33.3 2.6
14.3 32.1 2.3
14.9 34.5 2.3
17.0 35.6 2.1
16.1 36.8 2.3
13.2 47.2 3.6
12.7 41.0 3.2
15.3 33.0 2.2
14.7 34.8 2.4
“Not identified.
sition of membrane lipids is significantly affected by temperature (Cossins et al, 1978) and that higher levels of short-chain PUFA occur at higher environmental temperatures (Martin and Ceccaldi, 1977). P. mono&n are farmed in shallow ponds ( - 1 m) with fluctuating and often high temperatures and possibly 18 : 2n-6 may be utilized in preference to 20 : 4~6. Extensive samples (numblers 16, 26 and 28), however, although showing considerably higher levels of 18 : 2n-6 than wild P. monodon, also show higher levels. of 20 : 4n-6 than intensively farmed P. monodon. It would appear, therefore, that 18 : 2n6 is being substituted at least partly for 20: 4n-6 due to dietary deficiency. The possibility that 20: 4n-6 is an essential fatty acid for P. monodon should, therefore, be considered. If this is the case then none of the diets provide sufficient 20 : 4n-6 and widely quoted n-3/n-6 ratios may not be useful. The fatty acid profiles of the phospholipids of hatchery post-larvae reflect
C.D. O’LEARY AND A.D. MATTHEWS
16 TABLE 11 Fatty acid composition Fatty acid
of the phospholipids
Fatty acid composition 14 0.1
in samples of farmed P. monodon head
(O/o)
17
19
21
23
25
27
29
31
33
35
37
Av
14:o 15:o 16:0 16: 1 17:o 17: 1 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9 20: 1 n-7 20:2 20:4 n-6 20: 5 n-3 22:l n-11 2215 n-3 22: 6 n-3 24:l Others”
0.5 15.8 3.9 2.2 0.9 13.5 4.9 4.2 0.0 8.3 1.0 0.0 0.0 0.0 0.0 7.0 25.6 0.6 0.8 9.0 1.1 0.0
0.0 1.1 17.0 1.4 2.6 1.0 10.3 8.6 1.5 0.0 9.2 0.0 0.0 1.4 0.9 1.0 6.3 17.0 0.0 0.0 17.5 3.2 0.0
0.8 0.6 20.2 1.8 1.4 0.5 9.5 9.9 1.2 0.0 12.7 1.1 0.7 1.9 0.0 1.1 4.3 13.0 0.6 0.8 12.1 2.5 3.3
0.0 1.7 13.6 1.5 3.6 0.0 1.9 9.1 0.9 0.0 4.9 0.0 1.9 1.2 0.9 1.6 3.9 12.2 0.8 0.9 12.2 13.4 5.8
0.5 0.5 18.8 2.0 1.3 0.6 9.0 12.5 1.3 0.0 9.8 0.6 0.0 1.9 0.0 1.1 4.4 14.8 0.0 0.6 14.8 3.1 2.4
0.9 0.9 8.9 2.4 2.7 2.3 9.5 8.3 1.4 0.0 4.9 0.7 0.0 0.8 0.6 0.6 10.4 17.8 0.0 1.5 14.1 3.1 7.6
0.6 0.9 18.5 3.1 2.0 1.2 10.9 9.0 2.1 0.0 7.7 0.6 0.0 0.8 0.4 0.7 8.5 17.0 0.0 0.8 12.8 0.4 2.0
1.4 0.0 25.7 2.7 0.8 0.5 8.5 21.3 0.0 0.0 12.5 0.6 0.0 2.7 0.0 0.8 2.2 8.7 1.6 0.0 8.8 1.0 0.2
0.6 0.3 23.3 2.0 1.2 0.3 10.3 11.7 0.6 0.0 11.7 0.5 0.0 0.3 1.4 0.7 4.4 14.2 0.4 0.5 13.9 0.6 1.1
0.6 0.3 24.1 1.8 1.1 0.4 9.7 12.2 1.5 0.3 10.9 0.4 0.0 2.2 0.0 0.9 2.8 13.3 0.6 1.0 14.7 0.5 0.7
0.0 0.0 25.0 2.8 0.0 0.0 9.3 14.9 1.7 0.0 8.3 0.0 0.0 4.1 0.0 0.0 3.5 11.6 1.7 0.0 14.2 2.2 0.1
0.3 0.3 19.2 1.8 1.1 0.4 11.7 13.0 0.1 0.0 9.6 0.3 0.0 0.0 1.1 1.0 6.0 15.9 0.3 0.9 14.5 0.8 1.1
0.6 0.5 19.7 2.3 1.5 0.7 10.2 11.5 1.5 9.6 0.5 1.5 0.4 0.7 5.4 15.4 0.6 0.6 13.3 1.7 1.8
n-6 PUFA n-3 PUFA n-3/n-6
15.3 36.4 2.4
15.5 34.5 2.2
17.0 27.7 1.6
8.8 21.2 3.1
14.2 30.8 2.2
15.3 34.1 2.2
16.2 31.2 1.9
14.7 18.1 1.2
16.1 29.1 1.8
13.7 29.4 2.2
11.8 29.8 2.2
15.6 31.6 2.0
15.0 29.8 2.0
“Not identified.
their diets. Diatoms (Skeletonemu costatum, Chaetoceros, etc. ) have low levels of 22 : 6n-3 and high levels of 20: 5n-3 as have marine-type Artemia. Interestingly, the samples of klekap (benthic algae found in grow-out ponds), which forms part of the food of P. monodun in extensive and semi-intensive systems, also show high 20: 5n-3 and low 22 : 6n-3 levels. Millamena and Quinito ( 1985) have suggested that larval diets are generally deficient in 22 : 6n-3 and this analysis of wild and farmed post-larvae appears to confirm this. Of note is that the hatchery stock appear to have been fed freshwater-type Artemia because of the high level of 18 : 3s3. All the hatcheries had poor survival rates ( 1O-30% to PL20). Membrane fluidity may also be adjusted by altering the cholesterol content (higher cholesterol giving a more rigid structure, Deenan, 1965). The elevated levels of cholesterol in farmed P. monodon muscle may possibly be in
LIPIDS AND FATTY ACIDS IN PENAEUSMONODON
TABLE 12 Phospholipid
fatty acid composition
Fatty acid
Sample no. 39
14: 1
15:o
of post-larvae P. monodon
0.0 0.0
40
41
42
0.4
0.0
0.0
2.1 0.8 20.5 5.4 2.7 0.4 13.0 3.0 2.3 0.0 2.1 1.1 0.3 0.9 0.5 4.6 14.8 0.6 20.4 0.5 4.0 6.7 36.9 5.5
16:O 16:l 17:o 17:l 18:0 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2n-6 18:3 n-3 20: 1 n-9 20: 1 n-7 2012 2014 n-6 20: 5 n-3 2215 n-3 22: 6 n-3 24:l Others”
14.8 0.9 1.3 0.0 12.9 10.4 3.2 0.0 5.2 12.2 1.3 0.0 0.9 3.4 17.8 0.6 6.2 5.5 3.4
15.3 0.8 1.6 0.5 12.5 8.7 3.3 0.0 6.4 19.0 0.9 0.0 0.8 5.2 17.0 0.8 3.0 2.6 4.9
0.0 15.8 0.8 1.5 0.7 11.6 11.1 3.5 0.0 7.1 15.2 1.1 0.0 1.0 2.7 13.6 0.4 2.9 4.7 6.3
n-6 PUFA n-3 PUFA n-3/n-6
8.6 36.8 4.3
11.6 39.8 3.4
9.8 32.1 3.3
“Not ident:,fied.
response to elevated temperatures or because inadequate phospholipid of the correct fatty acid composition was available. Kanazawa et al. ( 1985 ) have shown that phosphatidylcholine and phosphatidylinostol containing the unsaturated fatty acids 20: 5n-3 and 22 : 6n-3 were effective in improving growth and survival of P. juponicus larvae. They considered that synthesis of these two phospholipids by penaeids is too slow for their needs and dietary supplementation using lecithin is essential. Analysis of data from Kanazawa et al. ( 1979b, 1985 ), Deshimaru et al. ( 1985 ) and Teshima et al. ( 1986 ) indicates a total dietary requirement for phosphatidylcholine and phosphatidylinositol containing 20 : 5n-3 and 22 : 6n-3 in the a position of 0.32 to 0.45% in the diet. Various commercial soyabean lecithins are available containing phosphatidylcholine (PC) from 25-97%, as are concentrates of phosphatidylinositol (PI) and therefore, the lecithin formulation can be manipulated. These results indicate that the lecithin source used should
16.7 29.7 1.8
4.6 0.6 20.1 4.0 0.4 0.0 3.2 9.0 1.1 0.3 15.7 2.2 2.2 3.3 1.0 0.7 11.4 2.1 1.3 11.9 0.4 4.5
14.8 17.8 1.2
4.7 0.1 21.3 4.7 0.6 0.3 4.2 11.2 0.3 0.0 13.4 1.6 0.7 2.9 1.4 0.1 6.5 1.6 1.3 7.6 1.5 14.0 16.9 25.3 1.5
4.0 0.4 20.8 5.5 0.9 0.7 5.4 14.4 0.6 0.0 15.9 1.5 0.9 1.9 1.0 0.2 9.6 1.5 2.0 11.1 0.3 1.4
3.7-5.7 0.0-0.4 17.6-24.6 3.5-6.0 o.o- 1.o 0.0-0.8 3.4-5.3 6.6-14.4 0.0-1.1 11.6-15.3 1.2-1.8 o.o- 1.4 1.5-5.8 0.0-2.1 0.0-0.4 3.7-9.3 1.1-2.0 0.8-1.7 3.7-10.2 0.0-2.1
Average
Range
Tungpao”
11.1-18.9 0.9-1.8 0.8-1.0 1.5-2.1 0.8-1.2 0.0-0.5 7.5-l 1.3 1.2-1.9 1.7-2.7 8.1-14.5 0.0-0.6
11.2 26.5 2.4
9.5-9.6 3.2-4.2 1.2-1.3 11.4-12.7 0.6-0.7
4.0-4.9 0.8-1.3
9.3-l 1.2 1.1-1.3
17.7 23.3 1.3
12.2-19.2 1.5-2.6 1.2-1.8 3.6-4.2 0.8-1.3 0.6-1.1 6.1-10.3 1.8-2.8 1.4-2.1 7.0-l 1.4 0.7-2.1
3.8-4.9 0.0-0.8 19.5-21.4 3.8-4.7 o.o- 1.o o.o- 1.o 4.1-4.6 8.9-10.7 1.0-1s
4.3 0.3 20.3 4.1 0.7 0.4 4.3 9.9 1.2 0.0 16.7 2.1 1.4 3.9 1.0 0.8 7.7 2.3 1.7 9.6 1.2 6.1 5.7-6.4 0.5-0.6 22.7-23.8 5.4-5.8
6.1 0.6 23.3 5.6 0.5 0.0 3.6 8.2 1.4 0.0 10.3 1.2 1.7 4.5 0.9 0.6 9.6 3.7 1.3 12.1 0.7 4.1
3.6-4.2 0.0-0.5 18.4-23.1 5.0-5.8 0.0-1.3 0.5-1.4 5.2-5.8 13.3-15.4 0.4-0.7 3.5-3.7 7.9-8.4 1.3-1.5
Range
Average
Range
Average
Chuen Shin”
Range
Rajaudang”
diet numbers 2, 5; Bama 0, I, 2, 3, 4, 5; Tungpao IA, 2, 3, 4, 5; Rajaudang 4, 5; Chuen Shin 1, 2, 3, 3M; President
1.1-1.4
3.1-3.4 0.9-1.0 0.6-0.7 11.1-l 1.7
3.1-3.3 8.7-9.2 1.0-1.1
4.5-4.7 0.5-0.6 18.8-21.4 3.5-4.5 0.3-0.4
Average
Average
Range
Bama”
of the total lipid of diets
Fanni”
“Average ofFanni ‘Not identified.
n-3 PUFA n-3/n-6
n-6 PUFA
20:4 n-3 20: 5 n-3 22:1 n-11 22: 5 n-3 2216 n-3 24: 1 Other?
20:4 n-6
14:o 15:o 16:O 16: 1 17:o 17:l 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9
Fatty acid
Average fatty acid composition
TABLE 13
15.5-23.9 1.5-2.1 0.9-1.7 1.6-3.3 0.7-1.1 0.0-0.6 7.5-10.4 1.1-2.3 1.2-1.8 8.0-12.4 0.0-1.1
18.5-22.0 3.7-6.0 0.7-1.2 0.0-0.8 3.7-5.1 11.1-13.2 0.7-1.2
2.7-4.4
Range
IA, IB, 2, 3, 4, 5.
20.4 23.2 1.1
3.6 0.0 20.5 4.8 1.0 0.5 4.5 12.6 1.0 0.0 19.5 1.8 1.2 2.3 0.9 0.3 9.0 1.6 1.5 9.4 0.5 3.5
Average
President”
0.7 13.0 5.3 13.2 8.4 1.9 0.6 0.7 12.9 13.6 15.5
Muscle
0.9 16.0 5.6 12.3 10.1 1.8 0.4 0.7 12.8 12.5 12.8
Head
P. monodon” wild
2.1 20.5 5.4 13.0 3.0 2.1 1.1 0.3 4.6 14.8 20.4
P. monodona post-larvae, wild
2.4 15.4 6.9 6.5 9.0 2.0 0.4 7.9 3.3 13.1 7.6
male
1.8 16.1 8.3 6.2 11.3 1.5 0.5 5.4 3.3 12.7 10.6
female
P. japonicusb whole body
BPhospholipid, all other total lipid. bGuary, 1974. ‘Deshimaru et al., 1985. dSchauer et al., 1980. ‘Fujita et al., 1980. ‘Total lipid levels dry weight; sample 1,0.95%; sample 2,0.52%. *Ackman et al., 1968.
14:o 16:0 16:l 18:O 18: 1 n-9 18:2 n-6 18:3 n-3 20: 1 n-9 2014 n-6 20: 5 n-3 22:6 n-3
Fatty acid
1.5 21.3 4.4 10.5 10.0 1.0 1.2 12.2 3.9 4.0 6.6
Shortnecked clam”
1.5 15.2 10.4 3.2 29.1 6.8 6.4 0.4 0.0 13.6 0.0
Marinetype brine shrimpd nauplii, Italy
and diets
0.5 7.9 5.8 5.9 26.3 5.2 21.0 0.0 0.6 0.3 0.0
Freshwatertype brine shrimp’ nauplii
Comparison of the levels of the major fatty acids of Z? monodon, P. juponicus, dietary components
TABLE 14
4.3 20.9 4.8 4.4 11.5 15.8 1.7 2.8 1.0 12.2 9.1
Average for growout diets
32.7 6.9 21.0 0.1 0.3 1.1 0.3 0.0 0.0 13.8 1.7
Siceietonema costatums 2
3.7 19.6 8.3 1.9 2.7 4.5 1.6 1.4 0.0 13.8 3.1
1
9.6 21.7 13.5 1.4 1.4 2.3 6.9 1.6 2.2 10.4 0.0
Kiekap’ Sample no.
G 8 g 2
9 ;“h 5 2
;
E z 8 1 0 ZJ 2 z
80
CD. O’LEARY AND A.D. MATTHEWS
have 50% PC and 4% PI containing the required percentage of the fatty acids 20: 5n-3 and 22: 6n-3 as indicated above.
REFERENCES Ackman, R.J., Tocher, CF. and McLachlan, J., 1968. Marine phytoplankton fatty acids. J. Fish. Res. BoardCan., 25(8): 1603-1620. Chanmugan. P., Donovan, J., Wheeler, C.J. and Hwang, D.H., 1983. Differences in the lipid composition of fresh water prawn (Macrobrachium rosenbergii) and marine shrimp. Food Sci., 48: 1440-1442. Cossins, A.R., Christiansen, J. and Prosser, C.L., 1978. Adaptation of biological membranes to temperature. Biochim. Biophys. Acta, 5 I 1: 442-454. Deenan, L.L.M., 1965. Phospholipids and biomembranes. In: R.T. Holman (Editor), Progress in the Chemistry of Fats and Other Lipids, VIII ( 1). Pergamon Press, Oxford, pp. 12-I 4. Deshimaru, O., Kuroki, K., Mazid, M.A. and Kitamura, S., 1985. Nutritional quality of compounded diets for prawn, Penaeus monodon. Bull. Jpn. Sot. Sci. Fish., 51(6): 1037-1044. Fujita, S., Watanabe, T. and Kitajima, C., 1980. Nutritional quality of Artemia from different localities as a living feed for marine fish from the view point of essential fatty acids. In: G. Persoone, P. Sorgeloos, 0. Roels and E. Jaspers (Editors), The Brine Shrimp Artemia, Vol. 3. Universa Press, Wetteren, Belgium, pp. 277-289. Gopakumar, K. and Nair, M.R., 1975. Lipid composition of five species of Indian prawns. J. Sci. Food Agric., 26: 3 19-325. Guary, J., Kayama, M. and Murakami, Y., 1974. Lipid class distribution and fatty acid composition of prawn, Penaeusjaponicus Bate. Bull. Jpn. Sot. Sci. Fish., 40( 10): 1027-l 032. Kanazawa, A., 1985. Nutrition of penaeid prawns and shrimps. In: Proc. 1st. Int. Conf. Culture of Penaeid Prawn/Shrimp, Iloilo City, The Philippines, 1984. SEAFDEC Aquaculture Department, pp. 123-l 30. Kanazawa, A. and Teshima, S., 1977. Biosynthesis of fatty acids from acetate in the prawn, Penaeus japonicus. Mem. Fat. Fish., Kagoshima Univ., 26: 49-53. Kanazawa. A., Teshima, S. and Sasaki, M., 1979a. Requirement of prawn, Penaeus japonicus for essential fatty acids. Mem. Fat. Fish., Kagoshima Univ., 33 ( I): 63-7 1. Kanazawa, A., Teshima, S., Tokiwa, S., Endo, M. and Razek, F.A.A., 1979b. Effects of short necked clam phospholipids on the growth of prawn. Bull. Jpn. Sot. Sci. Fish., 45( 8): 961965. Kanazawa, A., Teshima, S. and Tokiwa, S., 1979~. Biosynthesis of fatty acids from palmitic acid in the prawn, Penaeus japonicus. Mem. Fat. Fish., Kagoshima Univ., 28: 17-20. Kanazawa, A., Teshima, S., Ono, K. and Chalayondeja, K., 1979d. Biosynthesis of fatty acids from acetate in the prawn, Penaeus monodon and Penaeus merguiensis. Mem. Fat. Fish., Kagoshima Univ., 28: 2 l-26. Kanazawa, A., Teshima, S. and Sakamoto, M., 1985. Effects of dietary lipids, fatty acids and phospholipids on growth and survival of prawn (Penaeus japonicus). Aquaculture, 50: 3949. Kayama, M., Hirata, M., Kanazawa, A., Tokiwa, S. and Saito, M., 1980, Essential fatty acids in the diet of prawn. III. Lipid metabolism and fatty acid composition. Bull. Jpn. Sot. Fish., 46(4): 483-488. Martin, B.J. and Ceccaldi, H.J., 1977. Influence de la temperature sur la composition en acides gras du muscle abdominal de Palaemon serratus. Biochem. Systemat. Ecol., 5: 15 l-l 54. Metcalfe, L.D. and Wang, C.N., 198 1. Rapid preparation of fatty acid methyl esters using organic base-catalysed transesterification. Chromat. Sci., 19: 530-535.
LIPIDS AND FATTY AClDS IN PENAElJS MONODON
81
Millamena, O.M. and Quinito, E.T., 1985. Lipids and essential fatty acids in the nutrition of Penaeus monodon larvae. In: Proc. 1st Int. Conf. Culture of Penaeid Prawn/Shrimp, Iloilo City, T:he Philippines, 1984. SEAFDEC Aquaculture Dept., C: 18 1. Pozo, R., Lavety, J. and Love, M.R., 1988. The role of dietary a-tocopherol (vitamin E) in stabilising the canthaxanthin and lipids of rainbow trout muscle. Aquaculture, 73: 165-175. Ramsey, L.L. and Patterson, WI., 1948. Separation and determination of the straight chain saturatled fatty acids C5--C 10 by partition chromatography. J. AOAC, 3 1 ( 1): 139-l 50. Schauer, P.S., Johns, D.M., Olney, C.E. and Simpson, K.L., 1980. International study on Artemia IX. Lipid level, energy content and fatty acid composition ofthe cysts and newly hatched nauplii from five geographical strains of Artemia. In: G. Persoone, P. Sorgeloos, 0. Roels and E. Jaspers (Editors), The Brine Shrimp Artemia, Vol. 3. Universa Press, Wetteren, Belgium, pp. 365-373. Teshima, :S., Kanazawa, A. and Kukuta, Y., 1986. Effects of dietary phospholipids on growth and body composition of the juvenile prawn. Bull. Jpn. Sot. Sci. Fish., 52( 1): 155-l 58.
Aquaculture, 89 (1990) 65-81 Elsevier Science Publishers B.V., Amsterdam
Lipid class distribution and fatty acid cornposition of wild and farmed prawn, Penaeus monodon ( Fabricius )
Cliona D. O’Leary” and Anthony D. Matthewsb aOverseasDevelopment Natural Resources Institute, Central Avenue, Chatham Maritime, Chatham, Kent, ME4 4TB (Great Britain) bBio,compatiblesLtd., Brunel Science Park, Kingston Lane, Uxbridge, Middlesex, UB8 3PQ (Great Britain) (Accepted 10 January 1990 )
ABSTRACT O’Leary, (CD. and Matthews, A.D., 1990.Lipid class distribution and fatty acid composition and farmed prawn, Penaeus monodon (Fabricius). Aquaculture, 89: 65-8 1.
of wild
Total lipid levels of the muscle of wild and farmed Penaeus monodon were similar (4.35% and 4.66% dry weight) whilst head lipid levels (hepatopancreas) were higher in farmed P. monodon (farmed !).36%, wild 7.07%). The lipids of the muscle of wild P. monodon are mainly composed of phospholipid (70.5%), triglyceride (2.9%) and cholesterol (19.84Oh). Head lipids have higher triglyceride levels (9.18%) and lower levels of phospholipid (56.7Oh). Farmed P. monodon muscle lipids had lower levels of phospholipid (57.0%) and higher levels of triglyceride (8.46%) and cholesterol (24.17%) in comparison to wild P. monodon. Head lipids also showed lower phospholipid (39.23%) and higher triglyceride (25.93%). The composition of the phospholipids of wild and farmed P. monodon was similar excepting spingomyelin. The major fatty acids of the muscle phospholipids of wild P. monodon were 16:O (13.0%), 16:l (5.3%), 18:O (13.2%), 18:ln-9 (8.4%) 18:2n-6 (1.9%), 20:4n-6 ( 12.9%), 20: 5n-3 (13.6%) and 22: 6n-3 (15.5%). Head phospholipids had almost the same profile. Farmed P. monodon phospholipid fatty acids were similar except that levels of 18 : 2n-6 ( 9.8%) were higher and 16: 1 ( 1.79%) and 20:4n-6 (4.9%) lower. The lipid composition of hatchery and wild post-larvae (approximately post-larvae day 20) was dissimilar. Wild post-larvae lipid was almost entirely phospholipid (68.7%) and cholesterol (27.8%) whilst hatchery stock showed lower phosphohpid levels and significant amounts of triglyceride (5.9-15.6%). Comparison of the fatty acids of the phospholipids showed that wild post-larvae had lower levels of 18: ln-9, 18: 2n-6 and higher levels of 16: 1 and 22:6n-3 (20.4% wild, 4.0% farmed). Analysis of diets fed to farmed P. monodon showed low levels of phospholipid and very high levels of free fatty acids, indicative of dietary lipid deterioration. Diets used for P. monodon are those which have been developed for P. japonicus. Differences, however, exist in the lipid composition and fatty acid profiles of the two wild stocks suggesting differing lipid formulations may be required in diets.
0044-8486/90/$03.50
0 1990 - Elsevier Science Publishers
B.V.
66
CD.
O’LEARY
AND
A.D.
MATTHEWS
INTRODUCTION
Culture of the tiger prawn, Penaeus monodon is increasing substantially in subtropical and tropical countries. Whilst many papers have been presented concerning the nutritional requirements of Penaeus juponicus, those for Penaeus monodon are sparse and specific dietary requirements for this species have not been established. Phospholipids have been shown to be essential in the diets of penaeids (Kanazawa et al., 1985; Teshima et al., 1986) as have the highly unsaturated fatty acids 20 : 5n-3 and 22 : 6n-3 (Kanazawa et al., 1979a). Whilst phospholipids are almost certainly involved in lipid and cholesterol transport in the prawn (Teshima et al., 1986) they are also the main structural component of cell membranes and correct formation of these membranes is necessary in order that the cells may function effectively. Many intensive commercial farms in Indonesia have reported stressed or weak stock (personal communications to A.D. Matthews) and whilst numerous factors may be involved, the overwhelming importance of dietary lipids and particularly phospholipids and the absence of basic lipid information for P. monodon, led to the present study. MATERIALS AND METHODS
Samples of wild and farmed P. monodon were collected from fishermen and farms along the coastline near Jepara, Central Java, Indonesia, during November 1988. Farmed samples, large wild P. monodon and hatchery and wild post-larvae samples were obtained live and transported on ice to the marine laboratory of the University of Diponegoro, Jepara, for immediate analysis. Small wild P. monodon (less than 30 g ) were obtained un-iced from the lishing boats. Diets were purchased from local suppliers. Post-larvae were analysed whole. For all other samples, heads were broken from the tail and tail muscle was extracted from the carapace before analysis. Lipids were extracted by the method of Pozo et al. ( 1988 ) . Lipids were separated into class by thin layer chromatography on high performance silica gel plates, layer thickness 200 pm, having a preabsorbent layer (Whatman) using methyl acetate, propan- l-01, chloroform, methanol, KC1 (25 : 25 : 10 : 9 : 0.25%) as the developing solvent for phospholipids and hexane, diethyl ether, acetic acid ( 75 : 23 : 2 ) as the developing solvent for non-polar materials. The developed plates were sprayed with 8% orthophosphoric/cupric spray and charred in an oven at 180°C for 10 min. Quantification of lipid fractions was by densitometry ( Joyce-Loebl Chromoscan 3 ) . Phospholipid fractions for gas-liquid chromatography (GLC) were isolated by silicic acid column chromatography (Ramsey and Patterson, 1948 ). Fatty acid methyl esters were prepared by the method of Metcalfe and Wang ( 198 1) and separated by GLC on a Supel-
LIPIDS AND I=ATTY ACIDS IN PENAEUS MONODON
61
cowax 10 fused silica capillary column. Fatty acids were identified by comparison with a polyunsaturated fatty acid standard, marine source (Supelco, U.K.). RESULTS
Total lipid and lipid class distribution Total muscle lipid levels (Tables 1 and 2) of wild and farmed P. monodon were similar, ranging from 3.8 to 4.7% dry weight wild (average 4.35%) and 3.5 to 5.4% farmed (average 4.66%). Head total lipid levels were higher in farmed (average 9.36%) than wild (average 7.07%). The lipid class distribution (Tables 3 and 4) shows this extra lipid is triglyceride. Wild muscle had a higher level of phospholipid (wild average 70.54%, farmed 57.0%) and lower triglyceride (wild average 2.9%, farmed 8.46%) and a lower level of chlolesterol (wild average 19.84%, farmed 24.17%). The increase in cholesterol with decreasing phospholipid concentrations may be significant. Postlarvae (Table 5 ) total body lipids showed a similar pattern. Total lipid levels of the diets (Table 6 ) were within the range generally utilized for penaeids of 6-l 0%. Phospholipid levels in the diets were low (in general N 1.25%). This may be due to the breakdown of phospholipid since free fatty acid levels were very high (average 24.1% in the lipid). Cholesterol levels in the diets fall within th.e optimal range, 0.5-1.0% (Kanazawa, 1985). TABLE 1 Moisture and total lipid levels of wild P. monodon Sample no.” -
Sex
Head Muscle
1 3 5 7 9b 10”
2 4 6 8
11 12 13 14
Number of prawns in sample 16 14 5 3 2 2
male female female male
1 1 1 1
Whole body Weight (g)
lo.oei 1.7 16.9 X!I 3.7 37.7 + 10.2 81.1 f 5.5 98.1 139.7 152.1 169.3
Length (cm) 11.5t0.6 13.7kO.9 17.0f 1.1 21.8+ 1.0
24.8 27.5 27.5 28.0
Moisture (O/o)
Lipid (%)
Head Muscle
Head ~-
77.7 77.6 75.6 74.1 74.1 72.1
78.5 77.3 76.7 76.1
74.6 73.6 73.2 74.6
Muscle
wwd dw ww
dw
1.6 1.8 1.8 1.7 1.6 1.9
0.9 1.0 1.0 0.9
4.2 4.4 4.3 3.8
1.2 1.1 1.2 1.2
4.7 4.2 4.5 4.7
7.2 8.0 7.4 6.8 6.2 6.8
‘Samples l-4 and 5- 14 were obtained from the fishing villages of Wedung and Rembang, respectively. bHeads from samples 11 and 14. “Heads from samples 12 and 13. ?vw= wet weight, dw = dry weight. “Results as shown are mean k s.d.
16 18 20 22 24 26 28 30 32 34 36 38
15 17 19 21 23 25 27 29 31 33 35 37
15 44 20 50 1 lb 27 9 15 20 25
I
Stocking density ( nlm2)
3 6 6 6 6 8 not known 12 10 12 16 18
Age’ (weeks)
none A,B E C C none none E B A,C D C
Dietd
28 10 12 8 7 10 11 4 7 16 7 5
Number of prawns in sample
1.8+0.8 2.3+ 1.0 2.8k2.1 3.5 f 0.9 3.6t0.7 9.6f2.5 1 l.Ok3.0 12.6f4.7 14.0f2.3 19.0f4.9 39.2t7.3 42.0 f 9.9
6.5* I.1 7.1+1.0 7.321.8 8.2+ 1.2 8.5f0.5 11.6+1.0 12.2& 1.1 12.4+ 1.5 12.8kO.7 14.6+ 1.2 18.0+_ 1.1 18.6+ 1.3
74.5 76.6 74.3 72.1 74.4 72.6 75.7 75.9
78.3 77.0 74.2 75.8 76.0 73.7 75.9 77.4 2.6 0.8 2.5 1.6 1.5 1.9 2.7 3.2 3.0 2.4 2.0
9.8 6.4 7.4 9.7 12.5 10.9 9.9 8.3
1.2 1.4 1.2 1.3 0.8 1.0 1.3 1.3 1.2 1.1 1.0
ww
Ww
dw
Muscle
Head
Muscle
Head
Weight (g)
Length (cm)
Lipid (%)
Moisture (%)
Whole body
5.5 3.5 3.9 5.4 5.4 4.6 4.6 4.4
dw
“Samples 15-18, 25, 26, 33-36 were obtained from The Brackishwater Aquaculture Development Centre, Jepara: samples 21, 22, 27, 28, 37 and 38 from private farms near the village of Bulak Baru; samples 19, 20, 23,24 and 30 from a private farm near the village of Bondo and samples 3 l-32 from a private farm, Semarang. bWild post-larvae, all others hatchery stock. ‘Age beyond PL20. dSee Table 6.
Muscle
Head
Sample no.a
Moisture and total lipid levels of farmed P. monodon
TABLE 2
69
LIPIDS AND FATTY ACIDS IN PENAECJS MONODON
TABLE 3 Percentage lipid class distribution
of wild P. monodon
Sample no.
PL
TG
CHOL
FFA
MG/DG
CE
Muscle 2 4 6 8 11 12 13 14 Average
62.2 62.3 69.7 79.5 72.1 78.0 70.7 69.8 70.54+ 5.89
0.0 1.4 2.8 0.0 4.5 0.0 4.2 10.3 2.9 k 3.29
19.0 19.7 19.3 18.8 21.5 21.8 19.7 18.9 19.84f 1.10
8.3 10.9 6.7 0.0 0.0 0.0 0.0 0.0 3.24f4.31
0.0 3.1 1.5 1.7 1.9 0.2 5.4 1.0 1.852 162
10.5 2.6 0.0 0.0 0.0 0.0 0.0 0.0
48.2 49.2 60.5 54.6 62.3 65.4 56.7f6.51
8.0 14.6 9.3 6.0 10.0 7.2 9.18k2.75
22.6 14.2 17.3 22.1 19.0 18.8 19.Ok2.84
17.6 17.6 8.3 9.1 8.2 6.7 11.25f4.55
0.0 4.4 4.6 8.2 0.5 1.9 3.27 k2.82
3.6 0.0 0.0 0.0 0.0 0.0
Head 1 3 5 7 9 10 Average
“PL=phospholipid; TG= triglyceride; CHOL=cholesterol; monoglyo:rides/diglycerides; CE = cholesterol esters.
FFA=free
fatty
acids;
MG/DG=
Composition of the phospholipids The composition of the phospolipid of wild and farmed P. monodon (Tables 7 and 8 ) in respect of phosphatidylcholine, phosphatidylserine, phosphatidylinositol and phosphatidylethanolamine were not found to be significantly different. Sphingomyelin levels were, however, higher in wild muscle and head phospholipids (wild muscle 15.0%, head 9.03%; farmed muscle 11.39%, head 7.36%). Fatty acid composition of the phospholipidsof P. monodon The major fatty acids of both wild and farmed muscle and head phospholipidwere foundto be 16:0, 16: 1, 18:0, 18: ln-9, 18:2n-6,20:4n-6,20:5n-3 and 22 : 6n-3 (Tables 9, 10 and 11). The only major differences were in levels of 16 : 1 (wild muscle average 5.3%, head 5.6%; farmed average 1.7%, 2.3%), 18.2n-6 (wild muscle 1.901’0, head 1.8%; farmed 9.8%, 9.6%) and 20 : 4n-6 (wild muscle 12.9%, head 12.8%; farmed 4.9%, 5.4%). The same fatty acids (Table 12) dominate the fatty acid profiles of wild and farmed post-larvae phospholipids but hatchery stock had higher levels of 18 : 2n-6, 18 : 3n-3 and a low level of 22: 6n-3. The fatty acid profiles for the range of diets made by any one manufacturer were found to be similar (Table 13 ). Differences between products in the
70
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 4 Percentage lipid class distribution
of farmed P. monodon
Sample no.
PL”
TG
CHOL
FFA
Muscle 16 18 20 22 24 26 28 30 32 34 36 38 Average
42.4 49.9 60.5 48.6 68.9 38.9 67.8 49.6 69.5 62.5 61.4 64.1 57.0f 10.18
20.8 24.0 0.0 6.7 0.0 22.3 2.7 6.9 0.0 11.6 6.5 0.0 8.46 f 8.78
22.0 16.2 36.8 20.9 31.0 20.0 23.7 24.6 30.5 17.3 17.3 29.8 24.17f6.25
10.6 7.0 0.0 14.0 0.0 9.7 0.0 11.1 0.0 5.1 5.4 0.0 5.24+ 5.00
4.2 2.9 2.7 9.8 0.1 9.1 5.8 7.8 0.0 3.5 9.4 6.1 5.1223.30
Head 15 17 19 21 23 25 27 29 31 33 35 37 Average
31.8 36.5 42.8 30.3 46.0 34.7 48.8 35.0 36.6 38.0 44.1 46.2 39.23 + 5.86
39.1 40.9 8.0 11.5 11.0 34.5 33.6 17.8 43.3 36.6 28.9 5.9 25.93 f 13.46
16.6 13.4 25.3 15.1 23.0 13.5 16.7 14.3 18.8 13.4 17.1 20.1 17.27k3.72
12.5 9.1 20.7 30.9 9.8 12.2 0.9 17.8 0.0 8.6 5.1 17.1 12.06 f 8.33
0.0 0.1 3.2 10.2 10.2 5.1 0.0 15.1 I.3 3.4 4.8 10.7 5.34t4.84
“For explanation of abbreviations
MG/DG
see Table 3.
TABLE 5 Lipid class distribution
of post-larvae of P. monodon
Sample no.
Origin
Diet
No. of prawns in sample
Postlarval day
PL”
TG
39
bPT Yayasan Dian Desa
Skeletonema, Artemia, Frippak
2000
20
54.6
10.3 35.1
40
bPT Alam Windu Tama
Skeletonemu, Taiwanese inert
1500
22
52.5
15.6 22.7
41
bPT Urang Baruna
Skeletonema, Taiwanese inert
1700
16
63.7
5.9 20.3
10.1
42
Benteng Portugis
Wild
2000
30”
68.7
0.4 27.8
3.1
“For explanation of abbreviations bHatchery. ‘Estimated.
see Table 3.
CHOL
MG/DG
0.0 9.2
71
LIPIDS AND FATTY ACIDS IN PENAEUS MONODON
TABLE 6 Moisture, total lipid and lipid class distribution Diet Name
Origin
Designation
A
Fanni
Taiwan
2 5
7.8 9.0
9.9 9.8
not done 12.5 40.6 11.3
20.4
9.5
5.7
B
Bama
Indonesia
0 1 2 3 4 5
7.1 9.9 9.3 9.8 12.0 10.2
8.8 9.4 7.2 7.2 7.2 5.2
17.5 17.8 14.4 14.1 14.2 18.7
39.1 7.6 38.4 9.6 35.3 9.6 36.7 10.1 32.0 9.4 31.5 9.3
18.8 18.1 21.0 22.6 22.0 21.6
11.9 11.8 15.1 12.7 19.2 15.9
5.1 4.3 4.6 3.8 3.2 3.0
C
Tungp ao
Taiwan
1A 2 3 4 5
10.8 10.6 10.9 11.5 10.4
9.5 8.1 8.1 7.0 7.5
11.8 11.3 15.2 15.8 16.9
34.9 9.8 41.9 6.3 36.5 13.3 28.1 16.1 30.0 15.5
28.0 12.7 25.4 9.3 22.3 11.4 27.4 10.5 23.1 11.3
2.8 5.8 1.3 2.1 3.2
D
Rajaudang
Indonesia
4 5
10.1 8.2
6.8 5.1
11.2 7.6
28.0 13.4 26.3 9.7
32.5 12.6 30.6 22.1
2.3 3.7
E
Chuen Shin Taiwan
1 2 3 3mths
10.7 11.0 8.4 11.0
7.4 8.0 6.7 6.0
18.5 42.6 7.3 15.1 47.5 8.3 not done 21.7 36.5 11.3
16.0 12.5 14.5 11.8 16.2 11.7
3.1 2.8
IA 1B 2 3 4 5
8.8 9.5 9.5 10.2 11.1 9.4
9.3 10.1 8.5 7.8 6.9 9.1
12.5 15.5 17.7 7.9 13.8 11.7
22.3 7.5 15.1 5.0 16.2 7.0 53.0 1.1 27.2 12.2 15.6 14.9
1.6 2.0 1.4 5.9 0.0 4.1
F
President Wanyng
Taiwan
aDry weighi. bFor explanation of abbreviations
Moisture (o/o)
of diets for post-larvae Total lipid” (Oh)
PLb (%)
TG
40.4 48.2 45.5 17.1 31.8 45.7
CHOL FFA MG/DG
15.7 14.2 12.2 15.0 15.0 8.0
CE
2.6
see Table 3.
range seem to be in pellet size only. Levels of 20 : 512-3and 22 : 6n-3 are at or above the level of 1% recommended by Kanazawa et al. ( 1979a). High levels of 18 : 2n-6 were found (presumably from soya lecithin) and low levels of 20 : 4~6. DISCUSSION
Total 1:ipidlevels of wild P. monodon muscle, average 1.1% wet weight (ww ), were similar to those reported for other penaeids (Guary et al., 1974; P. japnicus 1.2%; Gopakumar and Nair, 1975, P. indicus 1.3%; Chanmugan et al., 1983, P. aztecus 1.O%). Head lipid levels of wild P. monodon were also low,
72
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 7 Composition Sample no.
Muscle 2 4 6 8 11 12 13 14 Average
of the phospholipids Phospholipid”
of wild P. monodon
(%)
LysoPC
PC
PS
PI
3.0 1.5 0.0 0.9 0.0 0.0 0.0 0.0
43.0 44.8 43.3 44.0 47.1 44.4 46.2 45.9 44.84f
9.7 8.8 10.4 9.9 8.1 8.6 8.6 9.2 9.16f0.73
3.3 3.2 4.8 5.2 2.9 3.7 3.9 4.2 3.9kO.75
26.0 27.1 25.0 24.0 28.1 27.4 26.3 27.5 26.43 + 1.30
15.0 14.6 16.5 16.0 13.8 15.9 15.0 13.2 15.0f 1.05
2.8 2.2 2.8 3.8 2.8 2.9 2.88 + 0.47
27.0 25.4 27.5 26.4 27.6 28.4 27.05 20.9
8.4 11.0 9.8 9.4 7.8 7.8 9.03rt 1.16
1.36
PE
sphing
Head
1 3 5 7 9 10 Average
2.7 2.8 0.0 1.5 0.0 0.0
50.4 50.8 52.8 51.0 53.4 53.6 52.0& 1.3
a.7 7.8 7.1 7.9 8.4 7.3 7.87 + 0.56
“LysoPC = lysophosphatidylcholine; PC=phosphatidylcholine; PS=phosphatidylserine; phosphatidylinositol; PE=phosphatidylethanolamine; sphing=spingomyelin.
PI =
1.7% ww. Total body lipid levels of P. japonicus have been reported to be 2.8% ww with the hepatopancreas containing 10.5% lipid (Guary et al., 1974). It would appear, therefore, that hepatopancreas total lipid levels are considerably lower in P. monodon than in P. japonicus. Farmed and wild P. monodon had similar total muscle lipid contents but the lipid level of farmed P. monodon heads was higher, the extra lipid being triglyceride. Dietary imbalance in terms of the phospholipids : triglyceride ratio in the total lipid of the diets or carbohydrate: lipid ratios may be responsible. The lipid of wild P. monodon muscle and post-larvae was almost entirely phospholipid whilst both the muscle of farmed P. monodon and post-larvae from hatcheries showed reduced phospholipid and increased triglyceride. Wild P. monodon head lipid contained a significant amount of triglyceride but again phospholipid levels were lower in farmed P. monodon and triglyceride higher. Actual phospholipid concentrations were not signficantly different in the P. monodon heads (4.0% wild, 3.7% farmed) but were in the muscle. InsufIicient dietary phospholipid may have been the cause. The reported muscle lipid composition for P. japonicus (Guary et al., 1974) of 22.1% phospho-
LIPIDS AND F’ATTY ACIDS IN PENAEUSMONODON
73
TABLE 8 Composition Sample no.
Muscle 16 18 20 22 24 26 28 30 32 34 36 38 Average Head 15 17 19 21 23 25 27 29 31 33 35 37 Average
of the phospholipids Phospholipid”
of farmed P. monodon
(%)
LysoPC
PC
PS
PI
PE
sphing
0.0
45.1 48.0 50.4 52.6 49.7 40.8 47.2 46.2 51.0 50.8 48.0 42.7 47.71 f 3.39
8.3 9.0 1.2 1.9 6.7 12.1 8.1 10.1 4.1 7.3 8.5 8.9 8.23k 1.74
3.9 3.3 2.8 2.3 2.2 3.8 2.7 4.0 1.1 2.9 3.6 3.4 3.OkO.81
30.8 28.6 31.2 28.9 33.7 28.0 28.6 25.8 34.8 26.9 28.0 29.4 29.56 f 2.5
11.9 11.1 8.4 7.0 7.7 15.3 13.4 13.9 8.4 12.1 11.9 15.6 11.39k2.82
50.4 53.5 55.0 55.0 48.9 44.9 51.0 53.5 50.2 52.5 63.3 52.6 52.51 k4.22
6.6 7.2 6.9 5.2 8.9 11.4 9.2 8.5 8.6 9.3 8.2 7.9 8.16& 1.51
3.9 3.5 3.0 2.3 3.4 3.1 3.4 3.3 3.9 3.9 3.5 2.3 3.34f0.53
32.1 28.6 28.0 32.0 29.4 28.6 27.0 26.0 28.9 26.9 24.6 28.9 28.42 f 2.1
7.0 1.2 7.1 5.5 9.4 11.4 9.4 8.1 8.4 1.4 0.4 7.0 1.36 k 2.56
0.0 0.0
1.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
0.0 0.0 0.0 0.0 0.0 0.0
0.0 0.6 0.0 0.0 0.0 1.3
aFor explanation of abbreviations
see Table 7.
lipid, 4 1.5% sterols (cholesterol) and 12.2% triglyceride with a similar composition for ovary and hepatopancreas tissues is very different from the present results for P. monodon and may reflect differing dietary needs. The absence of de novo synthesis of linoleic ( 18 : 2n-6)) linolenic ( 18 : 3n3 ) , eicosapentaenoic (20 : 5n-3 ) and docosahexaenoic (22 : 6n-3 ). acids from acetate-‘“C or palmitic acid-14C has been demonstrated in P. japonicus (Kanazawa and Teshima, 1977; Kanazawa et al., 1979~) and P. monodon (Kanazawa et al., 1979d). Conversion of dietary linoleic acid to arachidonic acid and linolenic acid to eicosapentaenoic and docosahexaenoic acid has been observed but the conversion rates were slow (Kayama et al., 1980). Dietary
74
C.D. O’LEARY AND A.D. MATTHEWS
TABLE 9 Fatty acid composition
of the phospholipids
Fatty acid Fatty acid composition
in samples of wild P. monodon
(O/o)
Muscle
Head
2
4
6
8
11
12
13
14
Av
1
3
5
7
9
10
Av
14:o 15:o 16:0 16:l 17:o 17:l 18:0 18: 1 n-9 18: 1 n-7 l8:2 n-6 18:3 n-3 20: 1 n-9 20: 1 n-7 20:2 20:4n-6 20:5 n-3 22:l n-11 22: 5 n-3 22:6n-3 24:1 Othersa
1.0 1.3 15.4 4.4 3.0 1.4 13.9 6.3 1.3 2.1 0.7 0.0 0.0 0.7 11.2 16.7 0.0 1.7 14.8 0.0 4.1
0.9 1.3 14.2 4.0 3.2 1.5 13.8 6.4 1.0 2.0 0.6 0.9 0.4 0.8 11.9 15.9 0.0 1.8 14.5 0.4 4.5
0.6 0.7 11.4 4.1 2.7 1.1 14.8 6.9 0.8 2.1 0.6 0.7 0.0 0.8 14.7 14.0 0.7 1.6 16.2 0.4 5.1
1.4 1.0 16.7 8.8 2.1 1.8 10.9 11.6 0.6 2.0 0.7 1.0 0.7 0.0 11.0 11.9 0.6 1.2 11.9 0.6 3.5
0.5 0.9 11.8 4.7 3.2 1.8 12.9 7.5 0.8 1.7 0.6 0.9 0.0 1.0 13.4 12.6 0.6 1.8 16.8 0.7 5.8
0.0 0.6 8.8 3.7 2.6 1.8 14.8 10.1 0.5 2.0 0.5 0.5 0.0 0.8 15.3 11.2 0.4 1.2 18.7 0.6 5.9
0.6 0.7 12.6 6.4 2.8 1.9 12.3 9.1 0.6 1.8 0.5 0.8 0.4 0.7 13.3 12.5 0.6 1.2 15.4 0.5 5.3
0.5 0.8 12.8 5.9 2.9 2.3 11.9 9.4 0.4 1.7 0.5 0.8 0.4 0.7 12.7 13.7 0.4 1.1 15.4 0.6 5.1
0.7 0.9 13.0 5.3 2.8 1.7 13.2 8.4 0.8 1.9 0.6 0.7 0.2 0.7 12.9 13.6 0.4 1.5 15.5 0.5 4.7
1.3 1.4 17.6 4.8 3.0 1.8 13.8 8.6 1.1 2.0 0.0 1.0 0.0 0.9 11.6 14.4 0.0 1.2 12.6 0.0 2.9
1.1 1.4 16.3 4.4 3.2 1.9 14.3 8.5 0.9 2.1 0.6 0.0 0.0 1.0 12.2 13.5 0.0 1.3 12.0 0.5 4.8
0.7 1.0 15.1 4.9 2.6 1.4 12.6 9.3 0.7 1.9 0.5 1.2 0.0 1.0 13.9 12.5 0.7 1.2 14.1 0.0 4.7
1.0 0.8 15.7 6.9 1.9 1.6 11.5 11.7 0.6 1.6 0.4 1.3 0.0 0.8 12.7 12.7 0.7 1.0 12.5 0.0 4.6
0.8 1.2 16.3 6.3 2.5 2.3 10.4 11.3 0.4 1.6 0.4 0.4 1.2 1.1 12.4 11.3 0.4 0.9 12.7 0.0 6.1
0.7 0.9 14.8 6.3 2.4 2.0 11.6 11.4 0.0 1.8 0.4 0.3 1.2 1.1 14.1 10.8 0.5 1.5 12.6 0.0 5.6
0.9 1.1 16.0 5.6 2.6 1.8 12.4 10.1 0.6 1.8 0.4 0.7 0.4 1.0 12.8 12.5 0.4 1.2 12.8 0.1 4.8
n-6PUFA n-3PUFA n-3/n-6
13.3 13.9 16.8 13.0 15.1 17.3 15.1 14.4 14.8 13.6 14.3 15.8 14.3 14.0 15.9 14.6 33.9 32.8 32.4 25.7 31.8 31.6 29.6 30.7 31.2 28.2 27.4 28.3 26.6 25.3 25.3 26.9 2.5 2.4 1.9 2.0 2.1 1.8 2.0 2.1 2.1 2.1 1.9 1.8 1.9 1.8 1.6 1.8
“Not identified.
sources of 20: k-3 and 22: 6n-3 are considered essential (Kanazawa, 1984) but not 20 : 4~6. In Table 14 the levels of the major fatty acids of P. monodon wild, P. japonicus, dietary components and commercial diets are compared. The major differences found between wild P. monodon and P. japonicus were in the fatty acids 20: 4n-6 and 20: ln-9. This may be a reflection of differing synthesis abilities, diet or environment. The fatty acid composition of the phospholipid of wild and farmed P. monodon was similar except for lower levels of 16: 1 and 20: 4n-6 and higher levels of 18 : 2n-6 in farmed. Adequate amounts of the essential fatty acids 20 : k-3 and 22 : 6n-3 seem to be provided by the diets. Phospholipids are the major component of biomembranes and are characterised by having a high content of polyunsaturated fatty acids that confer fluidity to the membranes. It has been demonstrated that fatty acid compo-
75
LIPIDS AND FATTY ACIDS IN PENAEUSMONODON
TABLE 10 Fatty acid composition Fatty acid
of the phospholipids
Fatty acid composition
in samples of farmed P. monodon muscle
(O/o)
16
18
20
22
24
26
28
30
32
34
36
38
Av
14:o 15:o 16:0 16:l 17:o 17:l 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9 20: 1 n-7 2012 20:4 n-6 20: 5 n-3 22:l n-11 2215 n-3 2216 n-3 24:l Others=
0.0 0.6 15.5 3.1 2.4 0.9 11.7 3.7 4.5 0.0 7.8 1.1 0.0 0.7 0.0 0.5 6.5 26.4 0.0 1.2 10.7 0.8 1.9
0.0 0.9 16.4 1.2 2.7 0.9 7.9 7.7 1.4 0.0 9.0 0.0 0.0 1.1 0.0 0.0 5.9 17.7 0.0 1.1 21.0 4.0 1.1
0.0 0.6 23.5 1.8 1.5 6.0 7.5 10.1 1.5 0.0 13.9 0.6 0.0 1.7 0.0 1.1 2.7 12.2 0.7 1.3 13.5 3.8 1.4
0.0 0.9 17.2 1.6 2.3 0.9 7.9 8.9 1.3 0.0 9.4 0.0 0.0 0.9 0.0 0.9 4.5 15.6 0.5 1.6 17.9 4.1 3.6
0.0 0.7 19.7 1.5 1.3 1.1 7.1 9.6 1.8 0.0 9.4 0.8 0.0 1.8 0.0 1.3 3.4 13.8 0.0 1.1 17.6 5.5 2.3
0.8 2.2 17.9 3.4 3.8 2.9 10.8 7.3 1.5 0.0 5.7 1.0 0.0 1.5 0.0 0.0 8.6 18.7 0.0 1.0 11.4 1.2 0.3
0.6 0.7 17.1 2.4 2.2 1.1 11.0 7.8 2.2 0.0 7.5 0.6 0.0 0.5 0.4 0.7 7.4 17.3 0.0 1.0 15.6 0.6 3.3
0.0 0.0 19.2 1.2 1.1 0.0 10.1 9.6 1.0 0.0 13.7 0.5 0.5 1.5 1.1 0.0 3.3 14.8 0.6 1.0 18.8 2.0 0.0
0.2 0.0 21.1 0.3 1.3 0.0 10.4 9.4 0.6 0.2 11.6 0.4 0.0 0.8 0.0 0.7 4.5 16.8 0.5 0.8 18.8 0.6 1.0
0.0 0.0 15.1 0.8 1.1 0.4 10.5 6.8 1.1 0.0 9.4 0.4 0.0 1.1 0.0 .08 3.8 18.5 0.3 1.0 27.3 0.6 1.0
0.0 0.2 17.7 1.6 1.0 0.4 9.2 8.9 0.8 0.0 8.8 0.3 0.0 1.7 0.0 0.7 3.9 16.9 0.4 0.8 23.0 1.9 1.6
0.0 0.0 19.7 1.5 1.1 0.6 11.6 13.6 0.8 0.0 10.9 0.0 0.0 0.7 0.0 0.9 4.4 16.2 0.0 0.8 16.0 1.1 0.0
0.1 0.5 18.3 1.7 1.8 0.8 9.6 8.6 1.5 9.8 0.5 1.2 0.1 0.6 4.9 17.1 0.3 1.1 16.1 2.2 -
n-6 PUFA n-3 PUFA n-3/n-6
14.3 39.4 2.8
14.9 39.8 2.7
16.6 27.6 1.7
13.9 35.1 2.5
12.8 33.3 2.6
14.3 32.1 2.3
14.9 34.5 2.3
17.0 35.6 2.1
16.1 36.8 2.3
13.2 47.2 3.6
12.7 41.0 3.2
15.3 33.0 2.2
14.7 34.8 2.4
“Not identified.
sition of membrane lipids is significantly affected by temperature (Cossins et al, 1978) and that higher levels of short-chain PUFA occur at higher environmental temperatures (Martin and Ceccaldi, 1977). P. mono&n are farmed in shallow ponds ( - 1 m) with fluctuating and often high temperatures and possibly 18 : 2n-6 may be utilized in preference to 20 : 4~6. Extensive samples (numblers 16, 26 and 28), however, although showing considerably higher levels of 18 : 2n-6 than wild P. monodon, also show higher levels. of 20 : 4n-6 than intensively farmed P. monodon. It would appear, therefore, that 18 : 2n6 is being substituted at least partly for 20: 4n-6 due to dietary deficiency. The possibility that 20: 4n-6 is an essential fatty acid for P. monodon should, therefore, be considered. If this is the case then none of the diets provide sufficient 20 : 4n-6 and widely quoted n-3/n-6 ratios may not be useful. The fatty acid profiles of the phospholipids of hatchery post-larvae reflect
C.D. O’LEARY AND A.D. MATTHEWS
16 TABLE 11 Fatty acid composition Fatty acid
of the phospholipids
Fatty acid composition 14 0.1
in samples of farmed P. monodon head
(O/o)
17
19
21
23
25
27
29
31
33
35
37
Av
14:o 15:o 16:0 16: 1 17:o 17: 1 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9 20: 1 n-7 20:2 20:4 n-6 20: 5 n-3 22:l n-11 2215 n-3 22: 6 n-3 24:l Others”
0.5 15.8 3.9 2.2 0.9 13.5 4.9 4.2 0.0 8.3 1.0 0.0 0.0 0.0 0.0 7.0 25.6 0.6 0.8 9.0 1.1 0.0
0.0 1.1 17.0 1.4 2.6 1.0 10.3 8.6 1.5 0.0 9.2 0.0 0.0 1.4 0.9 1.0 6.3 17.0 0.0 0.0 17.5 3.2 0.0
0.8 0.6 20.2 1.8 1.4 0.5 9.5 9.9 1.2 0.0 12.7 1.1 0.7 1.9 0.0 1.1 4.3 13.0 0.6 0.8 12.1 2.5 3.3
0.0 1.7 13.6 1.5 3.6 0.0 1.9 9.1 0.9 0.0 4.9 0.0 1.9 1.2 0.9 1.6 3.9 12.2 0.8 0.9 12.2 13.4 5.8
0.5 0.5 18.8 2.0 1.3 0.6 9.0 12.5 1.3 0.0 9.8 0.6 0.0 1.9 0.0 1.1 4.4 14.8 0.0 0.6 14.8 3.1 2.4
0.9 0.9 8.9 2.4 2.7 2.3 9.5 8.3 1.4 0.0 4.9 0.7 0.0 0.8 0.6 0.6 10.4 17.8 0.0 1.5 14.1 3.1 7.6
0.6 0.9 18.5 3.1 2.0 1.2 10.9 9.0 2.1 0.0 7.7 0.6 0.0 0.8 0.4 0.7 8.5 17.0 0.0 0.8 12.8 0.4 2.0
1.4 0.0 25.7 2.7 0.8 0.5 8.5 21.3 0.0 0.0 12.5 0.6 0.0 2.7 0.0 0.8 2.2 8.7 1.6 0.0 8.8 1.0 0.2
0.6 0.3 23.3 2.0 1.2 0.3 10.3 11.7 0.6 0.0 11.7 0.5 0.0 0.3 1.4 0.7 4.4 14.2 0.4 0.5 13.9 0.6 1.1
0.6 0.3 24.1 1.8 1.1 0.4 9.7 12.2 1.5 0.3 10.9 0.4 0.0 2.2 0.0 0.9 2.8 13.3 0.6 1.0 14.7 0.5 0.7
0.0 0.0 25.0 2.8 0.0 0.0 9.3 14.9 1.7 0.0 8.3 0.0 0.0 4.1 0.0 0.0 3.5 11.6 1.7 0.0 14.2 2.2 0.1
0.3 0.3 19.2 1.8 1.1 0.4 11.7 13.0 0.1 0.0 9.6 0.3 0.0 0.0 1.1 1.0 6.0 15.9 0.3 0.9 14.5 0.8 1.1
0.6 0.5 19.7 2.3 1.5 0.7 10.2 11.5 1.5 9.6 0.5 1.5 0.4 0.7 5.4 15.4 0.6 0.6 13.3 1.7 1.8
n-6 PUFA n-3 PUFA n-3/n-6
15.3 36.4 2.4
15.5 34.5 2.2
17.0 27.7 1.6
8.8 21.2 3.1
14.2 30.8 2.2
15.3 34.1 2.2
16.2 31.2 1.9
14.7 18.1 1.2
16.1 29.1 1.8
13.7 29.4 2.2
11.8 29.8 2.2
15.6 31.6 2.0
15.0 29.8 2.0
“Not identified.
their diets. Diatoms (Skeletonemu costatum, Chaetoceros, etc. ) have low levels of 22 : 6n-3 and high levels of 20: 5n-3 as have marine-type Artemia. Interestingly, the samples of klekap (benthic algae found in grow-out ponds), which forms part of the food of P. monodun in extensive and semi-intensive systems, also show high 20: 5n-3 and low 22 : 6n-3 levels. Millamena and Quinito ( 1985) have suggested that larval diets are generally deficient in 22 : 6n-3 and this analysis of wild and farmed post-larvae appears to confirm this. Of note is that the hatchery stock appear to have been fed freshwater-type Artemia because of the high level of 18 : 3s3. All the hatcheries had poor survival rates ( 1O-30% to PL20). Membrane fluidity may also be adjusted by altering the cholesterol content (higher cholesterol giving a more rigid structure, Deenan, 1965). The elevated levels of cholesterol in farmed P. monodon muscle may possibly be in
LIPIDS AND FATTY ACIDS IN PENAEUSMONODON
TABLE 12 Phospholipid
fatty acid composition
Fatty acid
Sample no. 39
14: 1
15:o
of post-larvae P. monodon
0.0 0.0
40
41
42
0.4
0.0
0.0
2.1 0.8 20.5 5.4 2.7 0.4 13.0 3.0 2.3 0.0 2.1 1.1 0.3 0.9 0.5 4.6 14.8 0.6 20.4 0.5 4.0 6.7 36.9 5.5
16:O 16:l 17:o 17:l 18:0 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2n-6 18:3 n-3 20: 1 n-9 20: 1 n-7 2012 2014 n-6 20: 5 n-3 2215 n-3 22: 6 n-3 24:l Others”
14.8 0.9 1.3 0.0 12.9 10.4 3.2 0.0 5.2 12.2 1.3 0.0 0.9 3.4 17.8 0.6 6.2 5.5 3.4
15.3 0.8 1.6 0.5 12.5 8.7 3.3 0.0 6.4 19.0 0.9 0.0 0.8 5.2 17.0 0.8 3.0 2.6 4.9
0.0 15.8 0.8 1.5 0.7 11.6 11.1 3.5 0.0 7.1 15.2 1.1 0.0 1.0 2.7 13.6 0.4 2.9 4.7 6.3
n-6 PUFA n-3 PUFA n-3/n-6
8.6 36.8 4.3
11.6 39.8 3.4
9.8 32.1 3.3
“Not ident:,fied.
response to elevated temperatures or because inadequate phospholipid of the correct fatty acid composition was available. Kanazawa et al. ( 1985 ) have shown that phosphatidylcholine and phosphatidylinostol containing the unsaturated fatty acids 20: 5n-3 and 22 : 6n-3 were effective in improving growth and survival of P. juponicus larvae. They considered that synthesis of these two phospholipids by penaeids is too slow for their needs and dietary supplementation using lecithin is essential. Analysis of data from Kanazawa et al. ( 1979b, 1985 ), Deshimaru et al. ( 1985 ) and Teshima et al. ( 1986 ) indicates a total dietary requirement for phosphatidylcholine and phosphatidylinositol containing 20 : 5n-3 and 22 : 6n-3 in the a position of 0.32 to 0.45% in the diet. Various commercial soyabean lecithins are available containing phosphatidylcholine (PC) from 25-97%, as are concentrates of phosphatidylinositol (PI) and therefore, the lecithin formulation can be manipulated. These results indicate that the lecithin source used should
16.7 29.7 1.8
4.6 0.6 20.1 4.0 0.4 0.0 3.2 9.0 1.1 0.3 15.7 2.2 2.2 3.3 1.0 0.7 11.4 2.1 1.3 11.9 0.4 4.5
14.8 17.8 1.2
4.7 0.1 21.3 4.7 0.6 0.3 4.2 11.2 0.3 0.0 13.4 1.6 0.7 2.9 1.4 0.1 6.5 1.6 1.3 7.6 1.5 14.0 16.9 25.3 1.5
4.0 0.4 20.8 5.5 0.9 0.7 5.4 14.4 0.6 0.0 15.9 1.5 0.9 1.9 1.0 0.2 9.6 1.5 2.0 11.1 0.3 1.4
3.7-5.7 0.0-0.4 17.6-24.6 3.5-6.0 o.o- 1.o 0.0-0.8 3.4-5.3 6.6-14.4 0.0-1.1 11.6-15.3 1.2-1.8 o.o- 1.4 1.5-5.8 0.0-2.1 0.0-0.4 3.7-9.3 1.1-2.0 0.8-1.7 3.7-10.2 0.0-2.1
Average
Range
Tungpao”
11.1-18.9 0.9-1.8 0.8-1.0 1.5-2.1 0.8-1.2 0.0-0.5 7.5-l 1.3 1.2-1.9 1.7-2.7 8.1-14.5 0.0-0.6
11.2 26.5 2.4
9.5-9.6 3.2-4.2 1.2-1.3 11.4-12.7 0.6-0.7
4.0-4.9 0.8-1.3
9.3-l 1.2 1.1-1.3
17.7 23.3 1.3
12.2-19.2 1.5-2.6 1.2-1.8 3.6-4.2 0.8-1.3 0.6-1.1 6.1-10.3 1.8-2.8 1.4-2.1 7.0-l 1.4 0.7-2.1
3.8-4.9 0.0-0.8 19.5-21.4 3.8-4.7 o.o- 1.o o.o- 1.o 4.1-4.6 8.9-10.7 1.0-1s
4.3 0.3 20.3 4.1 0.7 0.4 4.3 9.9 1.2 0.0 16.7 2.1 1.4 3.9 1.0 0.8 7.7 2.3 1.7 9.6 1.2 6.1 5.7-6.4 0.5-0.6 22.7-23.8 5.4-5.8
6.1 0.6 23.3 5.6 0.5 0.0 3.6 8.2 1.4 0.0 10.3 1.2 1.7 4.5 0.9 0.6 9.6 3.7 1.3 12.1 0.7 4.1
3.6-4.2 0.0-0.5 18.4-23.1 5.0-5.8 0.0-1.3 0.5-1.4 5.2-5.8 13.3-15.4 0.4-0.7 3.5-3.7 7.9-8.4 1.3-1.5
Range
Average
Range
Average
Chuen Shin”
Range
Rajaudang”
diet numbers 2, 5; Bama 0, I, 2, 3, 4, 5; Tungpao IA, 2, 3, 4, 5; Rajaudang 4, 5; Chuen Shin 1, 2, 3, 3M; President
1.1-1.4
3.1-3.4 0.9-1.0 0.6-0.7 11.1-l 1.7
3.1-3.3 8.7-9.2 1.0-1.1
4.5-4.7 0.5-0.6 18.8-21.4 3.5-4.5 0.3-0.4
Average
Average
Range
Bama”
of the total lipid of diets
Fanni”
“Average ofFanni ‘Not identified.
n-3 PUFA n-3/n-6
n-6 PUFA
20:4 n-3 20: 5 n-3 22:1 n-11 22: 5 n-3 2216 n-3 24: 1 Other?
20:4 n-6
14:o 15:o 16:O 16: 1 17:o 17:l 18:O 18: 1 n-9 18: 1 n-7 18: 1 n-5 18:2 n-6 18:3 n-3 18:4 n-3 20: 1 n-9
Fatty acid
Average fatty acid composition
TABLE 13
15.5-23.9 1.5-2.1 0.9-1.7 1.6-3.3 0.7-1.1 0.0-0.6 7.5-10.4 1.1-2.3 1.2-1.8 8.0-12.4 0.0-1.1
18.5-22.0 3.7-6.0 0.7-1.2 0.0-0.8 3.7-5.1 11.1-13.2 0.7-1.2
2.7-4.4
Range
IA, IB, 2, 3, 4, 5.
20.4 23.2 1.1
3.6 0.0 20.5 4.8 1.0 0.5 4.5 12.6 1.0 0.0 19.5 1.8 1.2 2.3 0.9 0.3 9.0 1.6 1.5 9.4 0.5 3.5
Average
President”
0.7 13.0 5.3 13.2 8.4 1.9 0.6 0.7 12.9 13.6 15.5
Muscle
0.9 16.0 5.6 12.3 10.1 1.8 0.4 0.7 12.8 12.5 12.8
Head
P. monodon” wild
2.1 20.5 5.4 13.0 3.0 2.1 1.1 0.3 4.6 14.8 20.4
P. monodona post-larvae, wild
2.4 15.4 6.9 6.5 9.0 2.0 0.4 7.9 3.3 13.1 7.6
male
1.8 16.1 8.3 6.2 11.3 1.5 0.5 5.4 3.3 12.7 10.6
female
P. japonicusb whole body
BPhospholipid, all other total lipid. bGuary, 1974. ‘Deshimaru et al., 1985. dSchauer et al., 1980. ‘Fujita et al., 1980. ‘Total lipid levels dry weight; sample 1,0.95%; sample 2,0.52%. *Ackman et al., 1968.
14:o 16:0 16:l 18:O 18: 1 n-9 18:2 n-6 18:3 n-3 20: 1 n-9 2014 n-6 20: 5 n-3 22:6 n-3
Fatty acid
1.5 21.3 4.4 10.5 10.0 1.0 1.2 12.2 3.9 4.0 6.6
Shortnecked clam”
1.5 15.2 10.4 3.2 29.1 6.8 6.4 0.4 0.0 13.6 0.0
Marinetype brine shrimpd nauplii, Italy
and diets
0.5 7.9 5.8 5.9 26.3 5.2 21.0 0.0 0.6 0.3 0.0
Freshwatertype brine shrimp’ nauplii
Comparison of the levels of the major fatty acids of Z? monodon, P. juponicus, dietary components
TABLE 14
4.3 20.9 4.8 4.4 11.5 15.8 1.7 2.8 1.0 12.2 9.1
Average for growout diets
32.7 6.9 21.0 0.1 0.3 1.1 0.3 0.0 0.0 13.8 1.7
Siceietonema costatums 2
3.7 19.6 8.3 1.9 2.7 4.5 1.6 1.4 0.0 13.8 3.1
1
9.6 21.7 13.5 1.4 1.4 2.3 6.9 1.6 2.2 10.4 0.0
Kiekap’ Sample no.
G 8 g 2
9 ;“h 5 2
;
E z 8 1 0 ZJ 2 z
80
CD. O’LEARY AND A.D. MATTHEWS
have 50% PC and 4% PI containing the required percentage of the fatty acids 20: 5n-3 and 22: 6n-3 as indicated above.
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81
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