kenji taxii1,* , manabu seoka1, nana ohara1, toshiro nasu2
TRANSCRIPT
Aquaculture Sci. 55(4), 579 - 585 (2007)
Dietary Utility of Chilean Fish Meal and Pollack Liver Oil
for Juvenile Pacific Bluefin Tuna
Kenji TAxiI1,* , Manabu SEOKA1, Nana OHARA1, Toshiro NASU2, Sell ODA2,Shlgeru MIYASHITA2, late Masaharu UKAWA3, Sadao SHIMENO4
and Hidetsuyo HosoKAwA4
Abstract: Dietary utility of Chilean fish meal (FM) and pollack liver oil (PLO) for juvenile Pacific
bluefin tuna Thunnus orientalis were investigated to develop their artificial diet. Diets 1, 2 and 3 were composed of 73, 63 and 53% FM and 0, 10 and 20% PLO, respectively. Control diet was raw
sand lance (SL) Ammodytes personatus, with 77.6% crude protein and 6.2% crude lipid. Duplicate
groups of juvenile tuna, weighing 1.68 g, were fed each diet until apparent satiety 6 times daily for 10 days. The diets and SL led low survival of tuna, due to flash collision against the tank walls. As compared with SL, diets 1-3 sustained lower daily intake and specific growth rate. Reversely, diets 1-3 led higher feed efficiency, protein efficiency ratio, apparent protein retention, final carcass
crude protein content, entero-somatic index, and gastral peptic and enteral tryptic enzyme activities. Final carcass lipid and hepato-somatic index of tuna fed diets 2 and 3 were higher than those
fed diet 1 and SL. These results suggest that FM in diet is a less digestible protein source for juvenile tuna, while the amino acid balance of FM sustains their requirements due to better protein
retention than SL.
Key words: Thunnus orientalis; Dietary fish meal; Digestion and absorption; Digestive enzyme activity
An artificial full-cycle culture of Pacific bluefin tuna Thunnus orientalis was initially achieved by the Fisheries Laboratory, Kinki University (Sawada et al. 2005) . In order to
promote tuna aquaculture industries and business, further studies should be required to solve various problems, such as mass larval and juvenile mortality resulted from surface floating and bottom sinking, trauma caused by flash collision against tank walls and underde-veloped artificial diet (Miyashita et al 1999) . It is noteworthy to mention that the experimental design of juvenile tuna is also limited by less number of larval tuna with an incomplete seed
ling production and mass mortality with the trauma as cited above. These offer inevitable
problems, in shortening a rearing period as well as in restricting a statistical analysis, as cited by Glencross et al (2002), Aguado et al. (2004) and Takii et al. (2006).
To date, there have been a few studies on nutrient requirement and artificial diets for
southern bluefin tuna Thunnus maccoyii (van Barneverd et al. 1997; Clarke et al. 1997; Carter
et al. 1998; Carter et al. 1999). Carter et al
(1999) reported southern bluefin tuna showed lower FM digestibility than Atlantic salmon
Salmo salar in in vitro digestion. In our previous study (Takii et al submitting), juvenile tuna showed lower apparent digestibility coefficients of protein, lipid and energy for a moist
pellet, by mixing of equal parts of sand lance
Received June 13, 2007: Accepted September 18, 2007.1 Fisheries Laboratory, Kinki University, Uragami, Wakayama 649-5145, Japan.
2 Fish Nursery Center, Kinki University, Shirahama, Wakayama 649-2211, Japan.3 Fisheries Research Institute
, Marubeni-Nissin Feed Co. Ltd., Atsumi, Aichi 441-3605, Japan.4 Faculty of Agriculture, Kochi University, Nankoku, Kochi 783-8502, Japan.* Corresponding author . Tel.: 81-735-58-0116; fax: 81-735-58-1246. Email: [email protected]
580 K. Takii, M. Seoka, N. Ohara, T Nasu, S. Oda, S. Miyashita, late M. Ukawa, S. Shimeno and H. Hosokawa
and an artificial diet included Chilean fish meal
(FM), than chub mackerel Scomber japonicus. This suggested that juvenile tuna could less digest and utilize dietary FM which was denatured by severe heating and drying processes. Therefore, the present study was designed to ascertain dietary FM utility as a main protein source, together with pollack liver oil (PLO) utility as a lipid source, in order to develop an artificial diet for juvenile tuna. The populariza-tion of the artificial diet will intensely contribute to the establishment and improvement of their
aquaculture-techniques, such as seedling production, sustainable rearing strategy and meat
quality enhancement.
Materials and Methods
Test diet
Diets 1, 2 and 3 were included 73% FM and
0% PLO, 63% FM and 10% PLO, and 53% FM
and 20% PLO, respectively. Other ingredi
ents, such as wheat gluten,ƒ¿-potato starch,
vitamin mixture, mineral mixture, sodium
carboxy methyl-cellulose and cellulose were
commonly included in all diets (Table 1). The
PLO was mixed with 10% (w/w) of 70% DHA
oil (Harimakasei Co. Ltd., Tokyo, Japan), in
advance. Vitamin and mineral mixtures were
referred by Halver (1957) . After mixing these
ingredients and equal part of tap water (v/w),
a moist type pellet with 1 mm in diameter
was prepared by a laboratory pellet machine.
Chopped sand lance (SL) Ammodytes personatus
was also given as a control. The diets and SL
were stored in a freezer at - 20•Ž until use.
The proximate composition of diets 1-3 and
SL is also shown in Table 1. The crude protein
and ash contents of diets 1-3 fell with decreas
ing in FM inclusion. The crude lipid content of
diets 1-3 rose with an increase in PLO inclu
sion. There were no differences among reduc
ing sugar contents of diets 1-3. Whereas, SL
had higher crude protein content but lower
crude lipid and ash contents than diets 1-3.
The gross energy of diets 1, 2, 3 and SL was
ranged from 19,300 to 23,300 kJ/kg, calculated
using heat combustion coefficients, 23.01 kJ/g
Table 1. Dietary formula (%), proximate composition (% dry basis) and calculated gross energy of diets 1-4
1 Pollack liver oil containing 10% of DHA oil.2 Halver (1957) to
protein, 38.07 kJ/g lipid and 17.15 kJ/g sugar (DE Saliva and Anderson 1995).
Fish and feeding trial
Feeding trial was conducted in the Fish
Nursery Center, Kinki University, Shirahama,
Wakayama, and juvenile tuna were obtained by
an artificial seedling production. The fertilized
eggs were obtained from the brood stock in net
pen after mating at near sun set, at the aqua-
culture site of the Fisheries Laboratory, Kinki
University, Oshima, Wakayama. Each 25-30
juvenile tuna at 35 days posthatch, weighing
1.68 g, after acclimatization with a commercial
diet (Marubeni-Nissin Feeds Co. Ltd., Tokyo,
Japan) for 15 days, introduced into a 1 m3 black
circular tank. The duplicate tanks were pre
pared for each diet which was given until appar
ent satiation, 6 times daily (7:00, 9:00, 11:00,
13:00,15:00 and 17:00), for 10 days. The tank
was fixed with an inner green polyvinyl-sheet
to reduce mortality by the flash collision and
covered with white sunscreen. The feeding trial
was conducted under a flow through system at
6 1/mmn filtered seawater with gentle aeration.
Water D0 and temperature were maintained at
94.6•}5.2% (mean•}SD, n=10) and 26.0•}0.8•Ž
(n=10), respectively.
Dietary Utility of FM and PLO for Bluefin Tuna 581
Assays
At the end of the feeding trial, fish were indi
vidually weighed to measure gain, daily feed
intake, feed efficiency, protein efficiency ratio
(PER) and apparent protein and lipid retentions.
Hepato-, gastro- and entero (including pyloric
ceaca)-somatic indices and carcass proximate
composition were determined from 5 individual
fish and 3 pooled fish per tank, respectively. The
peptic and tryptic enzyme activities were assayed
using pooled gastral organ and enteral organ
including the pyloric ceaca of 5 fish per tank.
The proximate composition of diets, sand
lance and initial and final carcass were assayed
by AOAC methods (AOAC 1984). Dietary
sugar was measured by the phenol-sulfuric acid
method (Hodge and Hofreiter 1962). The gastral
and enteral organs were homogenized with iced-
eionized water (1: 9) . Aliquot of the resultant
homogenate was obtained by centrifugation at
10,000•~g and stored at - 80•Ž until proteinase
activity assay. Peptic and tryptic enzyme activi
ties were measured by the casein-Foline method
cited previously (Takii et al 1986). The reaction
mixture was incubated at 30•Ž, for 15 min. A
unit of enzyme activity was defined as l ,ƒÊmole
tyrosine liberated from casein for a min in a reac
tion mixture. The activities were expressed per g
tissue and per 100g body weight as total activity.
Statistical analysis Data were subjected to one-way ANOVA to
evaluate treatment differences (P < 0.05). If this was significant, differences among means were identified by Duncan's multiple range test (P < 0.05).
Results
Growth performance and organ somatic indices Growth performance and organ-somatic indi-
ces of juvenile tuna fed the diets 1- 3 and SL for 10 days are shown in Table 2. No significant difference in survival was found among the dietary treatments, but diets 1-3 sustained slightly higher survival than SL. Diets 1-3 led lower specific growth rate and daily feeding rate than SL. Feed efficiency, PER, apparent protein
Table 2. Growth performance and digestive organ-somatic index of juvenile PBT fed diets 1-4 for 10 days
1 Specific growth rate=100•~(in final mean weight - In initial mean
weight)/days; feed efficiency=100•~weight gain/feed intake;
protein efficiency ratio=weight gain/protein intake; apparent
protein and lipid retention=100•~protein and lipid retained/protein
and lipid ingested. 2 Hepato -
, gastro- and entero-somatic indices=100•~hepato-, gastro-
and entero-weight/body weight. Enteral organ was included pyloric
ceaca in entero-somatic index.
3The different superscripts indicates statistical differences in a row
(P< 0.05).
retention and entero-somatic index of juvenile tuna fed diets 1-3 were significantly higher than SL. Diets 2 and 3 with higher lipid contents caused in higher apparent lipid retention and hepato-somatic indices of tuna than those fed
diet 1 and SL with lower lipid contents.
Carcass proximate composition and proteinase activities
Carcass proximate composition and gastral
peptic and enteral tryptic enzyme activities are shown in Table 3. The highest final carcass moisture was obtained in juvenile tuna fed diet 1. The highest final carcass crude lipid content was detected in tuna fed diet 3, followed by diets
2, SL and 1. No significant differences were found in crude ash contents among the dietary treatments.
Final gastral peptic enzyme activity expressed as per g tissue indicated no difference among the dietary treatments, while diets 1 and 2 induced higher total activity than SL. The highest final enteral tryptic activity expressed as per
g tissue was obtained in tuna fed diet 1, followed by diets 2, 3 and SL. Diets 1- 3 led significantly higher total tryptic activities than SL.
582 K. Takii, M. Seoka, N. Ohara, T Nasu, S. Oda, S. Miyashita, late M. Ukawa, S. Shimeno and H. Hosokawa
Discussion
We obligated to design the present study with duplicate and short 10-day rearing period, owing to the difficulty of their seedling production and mass mortality due to the trauma. Miyashita et al (2000) and Sawada et al (2005) indicated the mass mortality of juvenile tuna due to the flash collision to tank wall frequently occurred from 44-day to 74-day posthatch. In the present study, high mortalities of juvenile tuna fed diets 1-3 and SL by the trauma were also detected, irrespective of setting the inner
green polyvinyl-sheet in the tank. This inevitable mortality offered us the difficulty of statistical analysis among the dietary treatments.
Takii et al (2007 submitting) reported that
juvenile tuna revealed lower apparent protein, lipid and energy digestibilities for a moist diet mixing equal-parts of raw sand lance and artificial FM diet than juvenile chum mackerel Scomber japonicus. The FM is ordinarily processed under heavy steam-boiling and heat drying. Therefore, we hypothesized that
juvenile tuna had less potency to digest heat damaged FM protein with undesirable conformation changes, but not in chum mackerel. In the present study, juvenile tuna given diets 1-3 included 73 - 53% FM revealed lower final mean body weight, specific growth rate and daily feed intake than those given SL. Moreover, diets 1-3 induced higher entero-somatic index and
enteral tryptic enzyme activity than SL. These results intensely support the previous hypothesis that juvenile tuna can not digest efficiently the heat damaged FM in diets. Less digestible FM might require long time necessary for digestion which, in turn, lowered feed intake and weight
gain in juvenile tuna. Moreover, the physiological adaptation for less digestible FM might also induce the enteral organ enlargement and hyper tryptic synthesis and secretion from diffused
pancreas. Shimeno et al (1993) examined the adaptive diversity of digestive and absorptive processes in young yellowtail Seriola quinqueradiata for raw sand lance and FM diet. The sand lance
and FM diet led a similar evacuation pattern of total digesta in digestive organs of yellowtail. Although, enteral digesta fed FM diet rose faster than that fed on sand lance in the early time after feeding. Furthermore, FM diet induced higher entero-somatic index and enteral tryptic and amyloritic enzyme activities in yellowtail. Takii et al (2000, 2001a, 2001b) also reported that red sea bream, Pagrus major, tiger puffer, Fugu rubripes and yellowtail increased in tryptic enzyme activities of enteral organ and/or pyloric ceaca for FM diets supplemented with soybean trypsin inhibitors. Fish including a typical carnivorous tuna may have a common tendency which stimu-lates digestive organs enlargement and digestive enzyme secretion for less digestible diets.
However, feed efficiency, PER and apparent
protein retention of tuna fed diets 1-3 were higher than tuna fed SL diet. This may strongly
Table 3. Carcass proximate composition and proteinase activities of digestive organ in juvenile PBT fed diets 1-4 for 10 days
1 A unit was defined as 1ƒÊmole tyrosine liberated from substarte at 30•Ž for a min in the reaction mixture.
2 Enteral organ was included pyloric ceaca.
3 The different superscripts indicates statistical differences in a row (P< 0.05).
Dietary Utility of FM and PLO for Bluefin Tuna 583
support that the protein quality of FM has no
problems for juvenile tuna. After once less digestible FM diet was assimilated, its preferable amino acid balance would lead to promote high protein retention and feed efficiency in
juvenile tuna. Kaushik et al (2004), Fournier et al (2004) and Refstie et al (2005) demonstrated that European seabass, Dicentrarchus labrax, turbot (Psetta maxima) and Atlantic salmon
(Salmo salar) fed less digestible diets, substi-tuted large parts of dietary FM with plant protein sources, maintained relatively high feed efficiency and PER but low weight gain.
An increase in dietary lipid levels led higher apparent lipid retention and hepato-somatic index. Although statistically insignificant, diets 2 and 3 induced higher apparent protein retention than diet 1. A dietary protein-sparing effect by lipid was detected in various fishes, dentex, Dentex dentex (Tibaldi et al 1996; Company et al 1999; Slalli et al 2004) rainbow trout, Oncorhyncus mykiss (Lee and Putnam 1973; Takeuchi et al 1978), red sea bream, Pagrus major (Takeuchi et al 1991), tiger puffer, Takifugu rubripes (Takii et al 1995) and yellowtail (Takeda et al 1975). It is reasonable that the protein spearing effect is also functioned in juvenile tuna like other fish. Reversely, higher carcass lipid and hepato-somatic index induced by diets 2 and 3 may also indicate that dietary suitable level of crude lipid is less than 18% for juvenile tuna.
Protein to energy ratio (P/E ratio) of diets 1 - 3 and SL was 31.9, 25.4, 20.3 and 33.2 g/MJ,
respectively. In the previous study using sand lance (Takii et al 2005), we suggested that a dietary P/E ratio for maintaining body weight
was 38.2 g/MJ, calculated from 5.46 g protein/ kg/day and 143 kJ/kg/day. Sand lance could sustain juvenile tuna growth without serious troubles in both previous and present studies. These results reveal that the dietary PIE for juvenile tuna was in the range from 33-38
g/MJ. The dietary P/E has been reported in several fish, 30.6 g/MJ for Asian seabass Laees calcarifer (Catacutan and Coloso 1995), 25.9
g/MJ for Japanese seabass Lateolabrax japonicas (Al et al 2004), 23.7 - 26.6 g/MJ for sunshine bass Morone chrysops x M saxatilis (Webster et
al 1995; Keembiyehetty and Wilson 1998), 21.1-35.2 g/MJ for channel catfish Ictalurus punctatus
(Garling and Wilson 1976; Reis et al 1978) , 27.5 - 29.5 g/MJ for mutton snapper Lutjanus analis
(Watanabe et al 2001). Therefore, juvenile tuna, as a representative migratory and carnivorous fish, require much more protein for sustaining steep weight gain than other cultural fish.
The present study revealed that the juvenile tuna could not effectively utilize dietary FM, besides its preferable amino acid balance. It also indicates that a suitable dietary lipid level might
be less than 18% for juvenile tuna. Further studies are required to identify other useful and practical protein sources, necessary for developing an artificial tuna diet.
Acknowledgements
The authors would like to thank General
Director Prof Hidemi Kumai and other staff of
the Fisheries Laboratory, Kinki University, for
their valuable assistance, advice and encourage-
ment. This research was partially supported
by a Grant-in-Aid for Scientific Research (S)
14104007 from JSPS and 21st Century COE
Program•gCenter of Aquaculture Science
and Technology for Bluefin Tuna and Other
Cultivated Fish•hfrom the Ministry of Education,
Science, Sports and Culture of Japan.
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Carter, C. G., M. P. Bransden, R. J. van Barneveld and S. M.
Clarke (1999) Alternative methods for nutrition
research on the southern bluefin tuna, Thunnus mac
coyii: in vitro digestibility. Aquaculture, 179, 57-70.
Catacutan, M. R. and R. M. Coloso (1995) Effect of dietary
protein to energy ratios on growth, survival, and body
composition of juvenile Asian seabass, Lates calcarifer
Aquaculture, 131, 128-133.
Clarkee, S., A. Smart, R. van Barneveld and C. Carter
(1997) The development and optimization of manufac
tured feeds for farmed southern bluefin tuna. Austasia
Aquaculture, 11, 59-62.
Company, R., J. A. Calduch, J. Perez-Sanchez and J. Kaushik
(1999)Protein sparing effect of dietary lipids in
common dentex (Dentex dentex) : a comparative study
and protein utilization in juvenile sea bass, Dicentrachus
labrax. Aquaculture, 72,115-129.
DE Saliva, S. S. and T A. Anderson (1995) Energitics. In •gFish
Nutrition in Aquaculture•h Chapman & Hall, London,
pp.15-40.
Fournier, V., C. Huelvan and E. Desbruyeres (2004)
Incorporation of a mixture of plant feedstuffs as sub
stitute for fish meal in diets of juvenile turbot (Psetta
maxima). Aquaculture, 236, 451-465.
Garling, D. L. and R. P. Wilson (1976) Optimum dietary
protein to energy ratio for channel catfish figerlings,
lctalurus punctatus. J. Nutr., 106, 1368-1375.
Glencross, B., C. Carter, J. Gunn, R. von Barneveld, K.
Rough and S. Clarke (2002) Southern bluefin tuna,
Thunnus maccoyii. In •gNutrient Requirement and
Feeding of Finfish forAquaculture•h (ed. by C.R Webster,
C. Lim), CABI Publishing, Oxon, pp. 159-171.
Halver, J. E. (1957) Nutrition of salmonid fish-III. Water-
solible vitamin requirements of Chinook salmon. J.
Nutr., 62, 225-243.
Hodge, J. E. and B.T. Hofreiter (1962) Determination of
reducing sugars and carbohydrates. In •gMethods in
Carbohydrate Chemistry. Vol•h (ed. by R. L. Whistler,
ML. Wolfrom), Academic Press, New York, pp.
388-389.
Kaushik, S.J., D. Coves, G. Dutto and D. Blanc (2004)
Almost total replacement of fish meal by plant protein
sources in the diet of a marine teleost, the European
seabass, Dicentrarchus labranx. Aquaculture, 230,
391-404.
Keembiyehetty, C. N. and R. P. Wilson (1998) Effects of
water temperature on growth and nutrient utilization
of sunshine bass (Morone chrysops•~Morone saxatilis)
fed diets containing different energy/protein ratios.
Aquaculture, 166, 151-162.
Lee, D.J. and G. B. Putnam (1973) The response of rainbow
trout to varying protein/energy ratios in a test diet. J.
Nutr.,103, 916-922.
Miyashita, S., N. Hattori, Y. Sawada, Y. Ishibashi, H.
Nakatsukasa, T Okada, O. Murata and H. Kumai (1999)
Ontogenetic change on oxygen consumption of bluefin
tuna, Thunnus thynnus. Suisanzoshoku, 48, 475-488.
Miyashita, S., Y. Sawada, T Okada, O. Murata and H. Kumai
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ク ロマグ ロ稚 魚 に お け るチ リ魚 粉 の利 用性
滝井健二 ・瀬 岡 学 ・大原菜奈 ・那須敏郎 ・小 田誠二
宮 下 盛 ・宇川正 治 ・示野 貞夫 ・細川秀毅
クロマ グロ配合飼料 の開発 に先立ち,チ リ魚粉(FM)と スケ トウダラ肝油(PLO)の 利用性につ
いて検討 した。体重1.689の 稚魚 に,対 照のイカナゴ切餌 と段階的にFMお よびPLOを 変化 させて
調製 した飼料1～3で,平 均体重1.689の クロマグロ稚魚 を10日 間飼育 した ところ,飼 料 区では対照
区より摂餌率 と成長率は顕著に低 く,逆 に,飼 料効率,タ ンパ ク効率,消 化管重値,体 脂質含量,消
化管プロテアーゼ活性な どは高い傾向 にあった。以上の結果か ら,ク ロマグロ稚魚 はFMを 効率良
く消化 で きないことが示 され,飼 料開発 に当たってはFM消 化性の改善 を図るか,あ るい は他 の適
正なタンパ ク源 を検索する必要のあることが示唆された。