Energy and protein requirements for maximum growth and maintenance of bodyweight of yellowtail

FISHERIES SCIENCE 2000; 66: 884–893

INTRODUCTION

Soft-dry pellets (SDP), a form of dry pellets developedalmost a decade ago, have gained wide acceptanceamong aquaculturists of Japan.1 To derive the maximumbenefit from the diet in terms of its performance as wellas economic returns, it is important to establish an ade-quate feeding rate. Previous studies on the nutrientrequirements of yellowtail used raw fish and moist pellets(MP) and gave results and feeding rates which cannot beapplied for extruded pellets (EP) because of their widelydiffering nutrient contents.2 In addition, the commer-cially recommended feeding standards vary among feedmanufacturers because the digestible energy contents ofdiets are not uniform. To determine an adequate feedingrate employing SDP, greater knowledge of the energy andprotein requirements for yellowtail is required.

Fish size and water temperature are the major factors

among the various physiological and environmental influ-ences on feeding activity in fish.3 In this series of studies,the energy and protein requirements of yellowtail seriolaquinqueradiata were previously determined at variousculture sites with different environmental conditionsduring the winter season, when the feeding activity iscomparatively low.4,5 The present study was conducted todetermine the energy and protein requirements of yellow-tail under different water temperatures (natural condi-tions), using fish of different sizes and diets of fixed energycontents. Very little information is available on theenergy requirement of marine fish. Hence, the data fromthe present study supplement our previous observationsand offer a better assessment of the energy and proteinrequirements of yellowtail.4,5

MATERIALS AND METHODS

Experimental diets

The dietary ingredients and proximate composition arepresented in Table 1. The experimental diets contained

Original Article

Energy and protein requirements for maximum growth andmaintenance of bodyweight of yellowtail

Kanako WATANABE,1 Yoichi HARA,2 Kenjirou URA,2 Takeyoshi YADA,2 Viswanath KIRON,1Shuichi SATOH1 AND Takeshi WATANABE1,*

1Laboratory of Fish Nutrition, Tokyo University of Fisheries, Minato, Tokyo 108-8477 and 2NagasakiPrefectural Institute of Fisheries, Nagasaki,Nagasaki 851-2213, Japan

SUMMARY: Six feeding experiments were conducted to determine energy and protein require-ments of different sized yellowtail seriola quinqueradiata (8–280 g) for maximum growth and main-tenance of bodyweight under different water temperatures (21.7–27.1°C) in 500 L polycarbonatetanks. For each experiment, fish were fed to satiation or fed at levels between 0 and 80% of the sati-ation level twice a day. The experimental diets were extruded dry pellets with known digestible energy(DE) and protein (DP) values. The growth rate was highest for the satiation group and decreasedproportional to the feeding levels. The DE (kcal/kgBW per day)and DP (g/kgBW per day) require-ments for maximum growth, calculated on the basis of the daily feeding rate of the satiation group,were 225 and 21.7, 136 and 14.8, 125 and 11.2, 111 and 10.7, and 92 and 8.2, respectively, for fishwith an average initial bodyweight of 8, 63, 160, 237, and 280 g. The requirements for maintenanceof bodyweight, based on when the growth rate was zero for the above sizes, were 61 and 5.9, 15and 1.7, 15 and 1.3, 24 and 2.3, and 12 and 1.0, respectively. The requirements for maintenance ofbody energy content, estimated on the basis of the relationship between the DE intake and energyretention, were significantly higher than those for maintenance of bodyweight.

KEY WORDS: energy requirement, feeding level, maintenance of bodyweight, maximumgrowth, protein requirement, seriola quinqueradiata yellowtail.

*Corresponding author: Tel: 81-3-5463-2546. Fax: 81-3-5463-0553.Email: [email protected]

Received 11 November 1999. Accepted 24 April 2000.

Energy and protein requirements of yellowtail FISHERIES SCIENCE 885

Japan) until the start of the feeding experiment whenthey were divided into six (Exps 1, 3–5) or seven (Exp. 2)groups. The feeding experiments were carried out in 500 L polycarbonate tanks at the facilities of the NagasakiPrefectural Institute of Fisheries; the experimental condi-tions are shown in Table 2. Fish weighing between 8 and280 g on average were separated into groups of 15 to 30 fish each according to the fish size and kept in thetanks with a water flow rate regulated at 15–20 L/min.

The energy and protein requirements were deter-mined based on the relationship between growth rateand ration size. Therefore, feeding was carried out at dif-ferent levels, 100% satiation to 0% satiation (starva-tion). The levels were decided after feeding all groups tosatiation and assuming that the group with the highestfeed intake was consuming at the satiation level. Fromthe subsequent day, the proposed feeding rates (as calcu-lated on the basis of the satiation group) were assignedto the different groups. Fish were fed their correspond-ing feeding levels twice a day (at 09:00 and at 16:00 h)for about 25 days, during which the average water tem-perature ranged between 21.7 and 27.1°C.

Sample collection and analysis

Fish from the starvation group were sampled every weekfor analyzing the body composition. The fish from theother groups were starved for 24 h before handling. Theinitial and final bodyweights were measured and 10 fishfrom each group were sampled for whole body andanother five fish for liver and muscle proximate compo-sition analysis. All samples were immediately stored at–30°C until analysis. The frozen samples were cut intosmall blocks and homogenized using a super-fibrator(Hasagawa Co., Ltd., Shizuoka, Japan). Samples wereanalyzed for protein, lipid, ash, moisture and gross energyby methods described previously.4

61–67% fish meal as the main protein source and 9–13%feed oil as the lipid source and were prepared as pelletsin two sizes (diet 1, 2 mm; diets 2 and 3, 6 mm) by usinga large twin screw extruder.6 They contained 41.4–49.6%crude protein (CP), 16.7–20.5% crude lipid (CL), andhad a gross energy (GE) of 4.8–5.0 kcal/g. The digestibleenergy to digestible protein ratio for the three experi-ments (based on values of energy digestibility as 85% andprotein digestibility as 90%4) ranged between 91–112,the optimal value for yellowtail being around 100 asreported earlier.7,8

Experimental set-up

The yellowtail were all of wild origin and reared on acommercial SDP (Sakamoto Feed Co., Ltd., Chiba,

Table 1 Dietary ingredients and proximate composition (%)of the experimental soft-dry pellets

Ingredient Diet 1 Diet 2 Diet 3(2 mm) (6 mm) (6 mm)Exps 1,4 Exp. 2 Exps 3,5

Local fish meal (Sardine) 67 71 61Wheat flour 8 8 14a-Starch 5 5 5Mineral + vitamin mixture 7 7 7Feed oil (Pollock) 13 9 13

Proximate composition (%)As is basis.

Crude protein 45.6 49.6 41.4Crude lipid 20.5 16.7 19.5Gross energy (kcal/g) 5.0 4.8 4.9

Digestible protein (%) 41.0 44.6 37.3Digestible energy (kcal/g) 4.3 4.1 4.2DE/DP 103.6 91.4 111.8

Table 2 Experimental conditions for yellowtail

Exp. 1 Exp. 2 Exp. 3 Exp. 4 Exp. 5

Initial fish size (g) 8 63 160 237 280Stocking density (fish/500 L) 30 25 18 15 15Feeding period (days) 25 26 26 23 24Feeding levels1 (%) 100 100 100 100 100

80 70 70 80 7060 50 50 60 5040 30 30 40 3020 20 10 20 100 10 0 0 0

0Water temp. (°C)

Range 22.5–30.0 25.8–28.5 22.0–28.5 23.8–27.0 18.2–23.9Average 27.1 27.1 24.6 25.0 21.7

1 Feeding levels were determined based on satiation levels assumed as 100%.

886 FISHERIES SCIENCE K Watanabe et al.

RESULTS AND DISCUSSION

Growth and feeding performance

The growth performances of yellowtail for the experi-ments are presented in Table 3. In the experiment withthe small size-group (Exp. 1), fish fed to satiation grewfrom 8 to 69 g, the growth rate being 721% as comparedto fish fed 20% which had a growth rate of 158%. Feedgain ratios were superior for fish fed to 80 and 60% satiation levels. The starved fish showed a 17.9%

decrease in their bodyweight after 1 week. In Exp. 5,where a large-size group of fish was employed, fish fed tosatiation produced the best growth, from an initial body-weight of 278 g to a final bodyweight of 411 g, with the growth rate being 47.8% and feed gain ratio being1.36. The growth rate of the 10% feeding level group wasas low as 0.3%. In each of the experiments, growth rateand condition factors were highest for the satiationgroups and growth decreased with a decrease of feedinglevels. There were positive correlations between: (i) thedaily feeding rate (DFR) and specific growth rate (SGR)

Table 3 Growth performance of yellowtail fed soft-dry pellets

Feeding Average BW1 (g) Growth SGR2 DFR3 Feed gain NEp4 Conditionratio Initial Final rate (%) (%) (%) ratio factor

Exp. 1Satiation 8.4 69.0 721.4 8.3 5.30 0.86 113.7 15.180% 8.4 60.6 621.4 7.9 4.90 0.81 94.1 14.660% 8.4 45.6 442.9 6.8 4.50 0.81 83.9 14.040% 8.4 41.1 389.3 5.1 3.90 0.87 80.7 14.120% 8.4 21.7 158.3 3.1 3.00 1.03 50.0 12.30% (15 days)5 8.4 5.4 -35.7 -2.9 0 0 -22.1 9.4

Exp. 2Satiation 63.0 164.5 161.0 3.8 3.32 0.97 66.2 14.370% 63.6 145.2 128.2 3.3 2.62 0.87 48.7 14.450% 63.6 122.1 92.1 2.6 2.20 0.83 30.0 14.030% 63.6 92.7 45.8 1.5 1.60 1.12 12.2 13.520% 63.6 86.6 36.2 1.2 1.16 0.98 6.2 12.910% 62.7 73.6 17.3 0.6 0.74 1.20 -7.9 12.10% (28 days)5 63.6 57.7 -9.3 -0.6 0 0 -25.1 11.1

Exp. 3Satiation 158.9 389.9 145.3 3.5 3.00 0.92 62.2 17.870% 165.8 322.7 94.6 2.6 2.30 0.95 50.9 16.450% 165.9 293.3 76.8 2.2 1.80 0.83 38.6 15.730% 161.0 230.7 43.3 1.4 1.30 0.91 20.6 15.110% 166.7 185.0 11.0 0.4 0.50 1.16 -6.6 14.10% (26 days)5 165.7 157.1 -5.2 -0.7 0 0 -16.8 14.1



1 BW, bodyweight.2 SGR, specific growth rate.3 DFR, daily feeding rate.4 NEp, productive energy (kcal/kgBW per day).5 Number of days of starvation are shown in parentheses.


experiments, there were no marked differences in proteincontents of whole body, but energy content increasedand moisture decreased with increasing bodyweight,probably depending on the seasonal variations due towater temperature as previously reported.9,10

Figure 4 shows the changes in the average proximatecomposition of whole body of starved fish from all theexperiments. Lipid and protein contents decreased andmoisture content increased with time, leading to adecreased energy content. This decrease in lipid contentswas marked as observed in previous studies on yellowtailand other species, where the bodyweight decrease wasaccompanied by a decrease in protein and an even largerdecrease in lipid.11–14

Retention of energy and protein

Figure 5 shows the total percentage of energy and proteinretained in the whole body of fish from each experiment.There was no marked difference in the retentions amongthe groups in Exp. 1 when compared with the values atthe lower feeding levels in the other experiments. Thetotal percentage retention of energy and protein was zerowhen digestible energy (DE) intake was less than 50 kcal/kgBW per day. With the exception of Exps 1 and3, generally there was a gradual increase in energy reten-tion as DE intake increased. In all experiments, thehighest energy retention values did not go above 50%whereas the maximum protein retention values were dif-ferent for the various fish sizes. In Exps 2 and 3, theprotein retention reached a peak at DE intake of around

as shown in Fig. 1 (Exp. 1); and (ii) the digestible energyintake and SGR as shown in Fig. 2 (Exps 1–5). Produc-tive energy (NEp = energy retention; kcal/kgBW perday) was highest in fish fed to satiation and decreasedcorresponding to the levels. The productive energy ofstarved fish (feeding ratio, 0%) was considered to beapproximately equal to the net energy value for bodymaintenance and activity (NEm). Feed gain ratios weresuperior in fish fed at levels slightly lower than satiation.Condition factors increased and DFR and growth ratedecreased with increasing bodyweight. The DFR of otherexperiments seemed to be a little lower than the usualvalues obtained when fish were stocked in cages, proba-bly because the rearing tanks were kept indoors or dueto the different stocking densities and low swimmingactivity.

Analytical results

The proximate composition and energy contents of thewhole fish body from all experiments are shown in Fig. 3. The lipid content was not determined for fishweighing 8 and 237 g (Exps 1, 4). Generally, the higherfeeding levels produced the larger fish and the data revealthat the moisture content decreased with an increase infish size and feeding levels. In Exps 1 and 2 employingthe smaller-size group of fish, the protein, lipid andenergy contents increased with fish size. In the other

Fig. 1 Relationship between daily feeding rate and specificgrowth rate of yellowtail based on data from Exp. 1: f(x) = 2.16x- 3.05, r = 0.99. The corresponding equations for other experi-ments were: Exp. 2: f(x) = 1.35x - 0.48 (r = 0.99); Exp. 3: f(x) =1.35x - 0.44 (r = 0.99); Exp. 4: f(x) = 1.24x - 0.68 (r = 0.99);Exp. 5: f(x) = 0.89x - 0.23 (r = 0.99).

Fig. 2 Relationship between digestible energy intake and specific growth rate of yellowtail from the different experi-ments. (�), Exp. 1: 8 g; (�), Exp. 2: 63 g; (�), Exp. 3: 160 g;(�), Exp. 4: 237 g; (�), Exp. 5: 280 g.


100 kcal/kgBW per day while the values of Exp. 4 werestill increasing, even above 100 kcal/kgBW per day. It isprobable that although the fish from the latter experi-ment were feeding at satiation level, they had to expendmore energy in acquiring feed as the diets were smallerin size. The larger fish (Exp. 5) reached a peak value ataround 60 kcal/kgBW per day.

The daily energy and protein retentions increasedproportionally with an increase in DE intake for allexperiments (Figs 6,7). Figure 7 also shows that generallywhen the daily energy retention is zero, the DE intake islower for larger fish and higher for smaller fish, indicat-ing size-dependent differences. A similar pattern in DE

intake was also observed when the daily protein reten-tion was zero.

Energy and protein requirements

Energy and protein requirements for maximum growthwere calculated based on the daily feeding rate of thesatiation group, whereas the requirements for mainte-nance of bodyweight were based on the value when theSGR was zero (Fig. 1). As an example, considering thefish with an initial bodyweight of 8 g, when the DFR forthe satiation group was 5.30%, the corresponding DEintake was 225 kcal/kgBW per day and digestible protein(DP) intake was 21.7 g/kgBW per day. In contrast, at aDFR of 1.43% the growth rate was zero (the equation isshown in Fig. 1), and the DE and DP requirements were61 kcal and 5.9 g/kgBW per day, respectively.

Overall, the requirements of DE and DP for maxi-mum growth ranged between 92 to 225 kcal and 8.2 to21.7 g/kgBW per day, whereas that for maintenance ofbodyweight ranged between 11.6 to 60.8 kcal and 1.0 to 5.9 g/kgBW per day, respectively, for fish weighing8–280 g, at water temperatures of 18.2–31.0°C. Therequirements for maintenance of bodyweight werehigher in Exp. 4, because the fish may have consumedmore energy in their pursuit of the relatively smaller-sized pellets offered in these experiments. Figure 8 chartsthe relationship between the energy requirement formaximum growth or maintenance of bodyweight and fishsize. The equations were derived on the basis of this rela-

Fig. 3 Contents of (a) moisture, (b) protein, (c) lipid, and (d)energy of whole body of fish from the different experiments.(�), 8 g; (�), 63 g; (�), 160 g; (�), 237 g; (�), 280 g.

Fig. 4 Per cent change in the mean whole body proximatecomposition (n = 5) during starvation, relative to the initialvalue, recorded for each sampling day. (�), moisture; (�),lipid; (�), energy; (•), protein.


of data is necessary to improve the precision of the afore-mentioned relationships. The protein and energyrequirements for the maintenance of whole body energyand protein contents were estimated from the relation-ship between DE intake and energy and protein reten-tion data. Figure 6, based on data from Exp. 1, indicatesthat energy retention was zero at a DE intake of 40.0kcal/kgBW per day (DFR = 0.94%) and protein retentionwas zero at 56.2 kcal/kgBW per day (DFR = 1.32%).Experiments 2 to 5 are illustrated in Fig. 7.

The relationship between protein intake and proteinor energy retention used for calculation of the respectivemaintenance requirements, as indicated by the corre-

tionship indicating the DE requirement for maximumgrowth (DERmax) and for maintenance of bodyweight(DERmaint) were:

However, if the requirements are to be calculated by sub-stituting for fish sizes we find that these values are lowerthan those for fish cultured in net cages. Comparing thepatterns depicted in Fig. 8, it appears that the DE require-ment for maintenance of bodyweight could be moreprone to environmental influences. Further accrual

DER kcal kgBW per day, maint = =-355 0 620 55 2x r. . . DER kcal kgBW per day, max = =-760 0 910 34 2x r. . ;

Fig. 5 Relationship between digestible energy intake andtotal (a) energy and (b) protein retention of yellowtail fromdifferent experiments. Refer to Fig. 2 for explanation ofsymbols.

Fig. 6 Relationship between digestible energy intake anddaily (a) energy [f(x) = 0.58x - 22.3, r = 0.99] and (b) protein[f(x) = 0.07x - 3.98, r = 0.99] retention of yellowtail from Exp. 1.


sponding equations for each experiment is shown in Fig. 9. The protein requirements for Exp. 1 were 3.8 g/kgBW per day for the maintenance of whole bodyenergy and 5.4 g/kgBW per day for protein maintenance.The DE and DP requirements for the maintenance ofwhole body energy ranged from 19.0 to 40.8 kcal and 1.7to 4.5 g/kgBW per day whereas those for the maintenanceof whole body protein were 10.6–55.9 kcal and 1.0–

5.4 g/kgBW per day, respectively (Table 4). The require-ments for maintenance of body energy balance werehigher than those for the maintenance of protein exceptfor the fish which had an initial weight of 8 g. Therefore,it can be suggested that the protein requirement ofsmaller-sized fish was higher due to their faster growth.

Similar studies on rainbow trout Oncorhynchus mykissindicated that the fish (initial BW 1.4–23.7 g) required80–100 kcal energy and 5.5–13.1 g protein/kgBW per dayfor maximum growth and 13–27 kcal energy and 1.5–1.7g protein/kgBW per day for maintenance of body-weight.15 In the case of larger trout weighing 73.8 g, thesevalues were 60–70 kcal and 4.0–4.6 g/kgBW per day formaximum growth and 21 kcal and 1.4 g/kgBW per dayfor maintenance of bodyweight. These values were from one-third to half those of yellowtail, probablybecause of a lower optimal rearing water temperature and slower growth Cyprium carpio of rainbow trout. In thecase of carp (initial BW 3.9–5.6 g), they required130–172 kcal and 11.1–17.2 g protein/kgBW per day formaximum growth and 18–35 kcal and 1.6–3.5 g/kgBWper day for maintenance of bodyweight.16 Again, despitethe almost identical sizes, these values were higher in yellowtail. Carp weighing 243.2 g required 68 kcal and5.45 g protein/kgBW per day for maximum growth and22 kcal and 1.7 g protein/kgBW per day for maintenanceof bodyweight at water temperatures between 25–27°C.When compared to yellowtail, the values for maximumgrowth are lower while those for maintenance are higher. Gatlin et al.17 reported that the energy and pro-tein requirements of channel catfish Ictalurus punctatus

Fig. 7 Relationship between digestible energy intake anddaily (a) energy and (b) protein retention of yellowtail fromthe different experiments (Data from Exps 2–5). The follow-ing equations represent this relationship for energy retention:Exp. 2: f(x) = 0.68x - 27.6 (r = 0.99); Exp. 3: f(x) = 0.67x - 16.7(r = 0.99); Exp. 4: f(x) = 0.76x - 26.8 (r = 0.99); Exp. 5: f(x) =0.63x - 11.5 (r = 0.99). The corresponding equations forprotein retention are: Exp. 2: f(x) = 0.07x - 1.62 (r = 0.97); Exp. 3: f(x) = 0.05x - 0.53 (r = 0.99); Exp. 4: f(x) = 0.06x - 1.71 (r = 0.99); Exp. 5: f(x) = 0.05x - 0.87 (r = 0.99). Refer to Fig. 2for explanation of symbols.

Fig. 8 Relationship between fish size and energy requirementfor (�), maximum growth [f(x) = 760x-0.34] and (�), mainte-nance of bodyweight [f(x) = 355x-0.53] of yellowtail fed soft-drypellets.


for maximum growth and 28–93 kJ (7–22 kcal) and1.5–2.5 g/kgBW per day for maintenance of bodyweightat 25.3–25.6°C. Requirements for maximum growthwere similar to yellowtail but the requirements for main-tenance of bodyweight were only half.

These results suggest that protein and energy require-ments for maximum growth of yellowtail are generallyhigher than those of other fishes. This could be due totheir faster growth and higher swimming activity. Thegrowth rate in terms of SGR (% per day) was 8.3 for yel-lowtail of bodyweight 8.4 g on average (Exp. 1) as against4.1 for red sea bream Pagrus major weighing 2.36 g.19 For237 g yellowtail (Exp. 4) the SGR was 2.6 as against 0.80 for rainbow trout of 220 g.20 When comparisons aremade with freshwater species, it should be rememberedthat their activity and optimal temperature ranges aredifferent.

The energy requirement for the maintenance of bodyenergy and activity (NEm) are estimated through theenergy loss during starvation and the relationshipbetween DE intake and energy retention. Energy loss perday during starvation was considered as the energyrequirement for the maintenance of body energy andactivity and it was 22.1 kcal/kgBW per day in yellowtailweighing 8 g (Table 3). When derived from the rela-tionship between DE intake and energy retention (Fig.6), the energy value at zero point on the vertical axis was22.5 kcal/kgBW per day. Although both values aresimilar, it is better to use the latter which are extrapo-lated from the graph for calculation of energy partitionwhile the former values are based only on losses over thestarvation period.

Energy budgets

It is difficult to estimate non-fecal energy loss of yellow-tail because they do not feed on non-protein dry diets.Ruchimat et al.21 estimated the non-fecal energy loss ofyellowtail weighing 12.8 g at 2.3–3.5% of DE intake.They also calculated metabolizable energy (ME) byassuming the value of non-fecal energy loss to be 2–4%of DE intake and subtracting non-fecal energy loss fromDE intake. The heat increment (HI) of feeding wasdetermined by subtracting the NEm value from the MEvalue. The daily energy retention obtained by this analy-sis is equal to the productive energy (NEp) value. Thus,calculated energy partitions of fish fed to satiation in thepresent study are shown in Table 5 and Fig. 10. and itwas found that 39.4–44.8% of GE intake was used forNEp, 7.9–19.5% for NEm and 17.3–34.8% for HI.Energy partitioning at 100% ME intake is shown inparentheses (Fig. 10). There was no marked difference inNEp which was around 40% among the different sizedfish (Table 5). For fish reared at similar water tempera-tures (Exps 1, 2) the NEm was higher for the larger fish,while the HI was lower.

(initial BW 8.3–8.8 g) for maximum growth and main-tenance of bodyweight were 100 kcal and 8.8 g/kgBW per day and 15 kcal and 1.3 g/kgBW per day, respec-tively. These values are also lower than those for yellowtail. In another study, McGoogan and Gatlin18

reported that the energy and protein requirements of reddrum Sciaenops ocellatus (initial BW 3.4–5.5 g) were776–958 kJ (185–229 kcal) and 20–25 g/kgBW per day

Fig. 9 Relationship between digestible protein intake anddaily (a) energy and (b) protein retention of yellowtail. Thefollowing equations represent this relationship for energy reten-tion: Exp. 1: f(x) = 6.03x - 22.3 (r = 0.99); Exp. 2: f(x) = 6.23x -27.6 (r = 0.99); Exp. 3: f(x) = 7.51x - 16.7 (r = 0.99); Exp. 4: f(x)= 7.92x - 26.8 (r = 0.99); Exp. 5: f(x) = 7.07x - 11.5 (r = 0.99).The corresponding equations for protein retention are: Exp. 1:f(x) = 0.76x - 3.98 (r = 0.99); Exp. 2: f(x) = 0.63x - 1.62 (r =0.98); Exp. 3: f(x) = 0.57x - 0.53 (r = 0.99); Exp. 4: f(x) = 0.59x- 1.71 (r = 0.99); Exp. 5: f(x) = 0.51x - 0.87 (r = 0.99). Refer toFig. 2 for explanation of symbols.


Masumoto et al.22 estimated energy budgets for yel-lowtail by feeding 12.8 g fish at 23–26°C on moist pelletshaving 52% crude protein (d.b.) and 16% lipid (d.b.).The energy partitioning was 33% for NEp, 17% for NEmand 36% for HI. These values were similar to the valuesof fish having an initial weight of 8–63 g in the presentstudy, even though stocking densities and the types ofdiets were different between the two experiments. Choet al.23 calculated the energy budgets in rainbow trout byan indirect measurement of the DE value, determiningoxygen consumption during the feeding or starvationperiod and assuming that energy lost through the gillsand the urine accounts for 8% of the total energy intake.The diet was prepared using fish meal and soybean mealand had 36% digestible protein (d.b.) and 16% lipid(d.b.). When fed to rainbow trout of 19–66 g initial BWat 15°C, the energy distribution was 49% for NEp, 17%for NEm and 5% for HI of total energy intake. The valuesof NEp and NEm were similar to our present study but

HI was higher. Ohta and Watanabe estimated energybudgets of rainbow trout and carp.15,16 The energy parti-tioning of rainbow trout weighing 23.7 g and fed a fishmeal diet having 31.9% crude protein and 18.7% crudelipid at 14–16°C were 41.4% for NEp, 7.6% for NEm and36.1% for HI from the total energy intake. Although therequirements were considerably different between yel-lowtail and rainbow trout, the energy budget was similarin smaller fish (initial bodyweight 8 g in Exp. 1). Thevalues for carp weighing 3.9 g and fed a fish meal diethaving 39.7% crude protein (w.b.) and 10.9% lipid (w.b.)at 22–27°C were 24.1% for NEp, 12.6% for NEm and31.9% for HI of total energy intake. The NEp of carp waslower than yellowtail, indicating that energy utilizationfor growth was higher in yellowtail.

Further experiments are needed to obtain moredetailed data on energy requirements of yellowtail underdifferent environmental conditions for various fish sizesin order to establish a feeding standard.

Table 4 Energy and protein requirements of yellowtail fed soft-dry pellets

Experiment 1 2 3 4 5

Initial fish size (g) 8 63 160 237 280Water temperature (°C) 22.5–30.0 25.8–28.5 22.0–28.5 23.8–27.0 18.2–23.9

(Average) 27.1 27.1 24.6 25.0 21.7Maximum growth

DFR 5.30 3.32 3.00 2.60 2.20DE1 225.3 135.5 124.8 110.5 91.5DP2 21.7 14.8 11.2 10.7 8.2

Maintenance of bodyweightDFR 1.43 0.37 0.35 0.56 0.28DE1 60.8 15.1 14.6 23.8 11.6DP2 5.9 1.7 1.3 2.3 1.0

Maintenance of body energyDE1 39.5 40.8 25.4 35.5 19.0DP2 3.8 4.5 2.3 3.4 1.7

Maintenance of body proteinDE1 55.9 25.2 10.6 29.8 19.0DP2 5.4 2.8 1.0 2.9 1.7

1 Digestible energy (kcal/kgBW per day).2 Digestible protein (g/kgBW per day).

Table 5 Energy partitioning in yellowtail fed soft-dry pellets

Experiment Initial fish size Water temperature DFR1 GE2 NEp3/GE2 NEm4/GE2 HI5/GE2

(g) (°C) (av.) (%) (%) (%)

1 8 22.5–30.0 (27.1) 5.30 280.6 40.5 7.9 33.1–34.82 63 25.8–28.5 (27.1) 3.32 168.0 39.4 18.3 23.9–25.63 160 22.0–28.5 (24.6) 3.00 155.3 40.1 10.8 30.7–32.44 237 23.8–27.0 (25.0) 2.60 137.6 44.8 19.5 17.3–19.05 280 18.2–23.9 (21.7) 2.20 113.9 43.1 10.2 28.3–30.0

1 Daily feeding rate (%).2 Gross energy intake (kcal/kgBW per day).3 Productive energy (NEp).4 Net energy value for maintenance of body energy and activity (NEm) (kcal/kgBW per day).5 HI = DE - (NEp + NEm).


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Fig. 10 Partitioning of dietary energy by yellowtail fed to satiation. Numbers in parentheses are the percentage of themetabolizable energy.

Energy and protein requirements for maximum growth and maintenance of bodyweight of yellowtail

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