compensatory growth and body composition of

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Compensatory growth and body composition ofjuvenile black rockfish Sebastes schlegelifollowing

feed deprivation

Sung-Yong OH,1* Choong Hwan NOH,1 Rae-Seon KANG,1 Chong-Kwan KIM,1Sung Hwoan CHO2 AND Jae-Yoon JO3

1Marine Resources Research Department, Korea Ocean Research & Development Institute,Ansan P.O. Box 29, Seoul 425-600, 2Division of Marine Environment and BioScience, College ofOcean Science and Technology, Korea Maritime University, Busan 606-791, and 3Departmentof Aquaculture, Pukyong National University, Busan 608-737, Korea

ABSTRACT: Compensatory growth, feeding rate, feed efficiency and chemical composition ofjuvenile black rockfish (mean weight 1.43 g) were investigated for 35 days after a 14-day feeddeprivation treatment under four feeding conditions: one group continuously fed (control) and the

other three groups fasted for 5 days (F5), 10 days (F10) and 14 days (F14). All fasted fish were re-fedfrom day 15. Only F5 achieved the same body weight as the control, indicating that completecompensation occurred in F5. The specific growth rate (SGR) of F5 was the highest at day 21 andthen decreased thereafter, showing higher values than the control at days 21, 28 and 42. In contrast,although SGRs of F10 and F14 were higher than that of the control during the whole refeeding periodexcept day 21, they did not catch up the control in body mass, indicating that only partial compen-sation occurred in F10 and F14. The feeding rate (FR) of all groups except F14 changed in a patternsimilar to SGR (Spearmans rank correlation, rs > 0.9), suggesting that SGR varied depending on FR.Similar feeding efficiencies (FEs) were found in the four groups and they did not vary significantlyduring the whole refeeding period, suggesting that FE was not the factor affecting SGR. At day 14, theratios of lipid to lean body mass in F10 and F14 were lower than those in the control and F5, and therewas no difference between the control and F5. At day 49, however, only F14 showed a lower valuethan the other three groups, and there was no difference among the three groups. These results

indicate that juvenile black rockfish fasted for 514 days can exhibit compensatory growth afterrefeeding, but timing and degree vary depending on the duration of feed deprivation.

KEY WORDS: black rockfish, body composition, compensatory growth, feed deprivation,refeeding.

INTRODUCTION

Compensatory growth refers to a phase of rapid

growth after a period of feed deprivation in order torecover an original body weight or growthtrajectory.13 Many organisms exhibit faster growthduring recovery from total or partial food depriva-tion than they do during periods of continuousfood availability, and thereby achieve the samesize as conspecifics experiencing environmental

conditions that are more favorable (reviewed in Aliet al.1). This is also true for fish species. Studies onfish including both coldwater and warmwater

species have shown that compensation growthoccurs following a period of feed deprivation orrestricted feeding (reviewed in Zhu et al.4). Asshown in these studies,1,4 however, the degree ofcompensation growth occurring after feed depri-vation is highly variable depending on experimen-tal species and feeding protocols including lengthand intensity of deprivation. For example, juve-niles of hybrid sunfish (F1 hybrid of female greensunfish Lepomis cyanellus male bluegill L.macrochirus) that received repeated cycles of nofeeding and refeeding exhibit significantly faster

*Corresponding author: Tel: 82-31-400-7728.

Fax: 82-31-406-2882. Email: [email protected]

Received 1 August 2007. Accepted 29 November 2007.

FISHERIES SCIENCE 2008; 74: 846852

2008 Japanese Society of Fisheries Science doi:10.1111/j.1444-2906.2008.01598.x


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growth rates than the control (i.e. overcompensa-tion growth),3 but juveniles of yellow perch Perca

flavescens that experienced a similar feedingregime to the hybrid sunfish juveniles do notexhibit such overcompensation growth.5 In addi-tion, some fish (e.g. channel catfish6 and Atlantic

salmon7

) have been reported to exhibit completecompensation achieving the same body mass ofthe control, whereas others (e.g. Alaska yellowfinsole8 and hybrid tilapia9) show partial compensa-tion in which food-restricted fish show acceleratedgrowth, but do not achieve the same body mass ofthe control. Such inconsistency, therefore, suggeststhat compensatory growth of fish species shouldbe fully understood before seeking a general trend.

Black rockfish Sebastes schlegeli is one of themost popular and commercially important marineaquaculture fish species in Korea, but little atten-tion has been paid to feeding practice including

combinations of feed deprivation and refeeding. Asrevealed in many studies, compensatory growth infish has aroused interest because of its theoreticalinterest8,1012 and potential application as a man-agement tool in commercial fish production.3,9,13 Inthis context, we investigated the difference of com-pensatory growth, feeding rate and feed efficiencyin juvenile black rockfish subjected to variousperiods of feed deprivation. In addition, to gaininsight into the effects of feed deprivation andsubsequent refeeding on the body compositionof juveniles, we also investigated the temporalchange in chemical attributes such as protein,

lipid, energy, ash and moisture content, and theratio of lipid to lean body mass.

MATERIALS AND METHODS

Experimental fish and rearing system

Black rockfish juveniles were obtained from aprivate hatchery (Ocean Tech, Tongyeong, Korea),transferred to a recirculating system, and thenacclimated for 10 days before experiment. Duringacclimation, fish were fed with a commercial dry

pellet (E-hwa Feed Co., Busan, Korea) with crudeprotein 46.4% and crude lipid 6.3%, to apparentsatiation twice a day at 09:00 and 16:00 hours.

The experiment was conducted using 20 circularacrylic chambers (30 cm diameter, 30 cm waterdepth, 21 L) in a recirculating system with aerationin each chamber. The water in each chamber wasdrainedatthebottomtoa5040-Lbiofilterthrougha190-L foam separator and pumped back to eachchamber at a rate of 41.5 L/h. During the experi-ment, dissolved oxygen, temperature, salinity,ammonia and pH were monitored daily. Water

temperature was maintained at 25.5 0.5C. Pho-toperiod was controlledat a 12:12 h lightdarkcycle

with the diurnal cycle from 08:00 to 20:00 hoursusing a 32-W fluorescent tube. Salinity ranged 33.034.3, dissolved oxygen was greater than 6.7 mg/L,ammonia nitrogen was less than 0.3 mg/L and pH

ranged 7.98.2 throughout the experiment.

Experimental design and sampling

After the 10-day acclimation, 200 juveniles [initialbody weight 1.43 0.01 g (mean standarderror)] were randomly assigned to one of fourfeeding groups: (i) 5-day feed deprivation (F5) fromdays 1014 after the beginning of experiment; (ii)10-day deprivation (F10) from days 514; (iii)14-day deprivation (F14) from days 114; and (iv) acontrol regularly fed to satiation twice a day at

09:00 and 16:00 hours throughout the wholeexperiment (49 days). Each group included fivechambers (i.e. replicates) and 10 juveniles wereheld per replicate.

Prior to the experiment, the weight of the juve-niles contained in each group was measured tocheck if there were differences in the initial body

weight between the four groups. For this, fish werefasted for a day to evacuate the gut, anesthetized

with 2-phenoxyethanol (Sigma, St. Louis, MO,USA) at 150 mg/L, and measurements were taken.Before measuring, all excess water was removed

with blotting paper. Fishes were weighed to 0.01 g.

To assess the effect of the four feed deprivationtrials, weights of the juveniles contained in eachgroup were re-measured at the end of day 14, andthen the three feed-deprived groups were fed againfrom day 15 using the same method applied to thecontrol. While feeding, a small weighed quantity offood was dropped into each chamber at intervalsof 5 min until the fish stopped consuming food.Each feeding lasted approximately 2 h. To comparethe growth rate of the four groups, the weight of the

juveniles was measured weekly after day 14 usingthe same method mentioned above. Fish were notfed on the day of weighing.

Chemical composition

Chemical attributes such as protein, lipid, energy,ash and moisture content, and the ratio of lipid tolean body mass (lipid/LBM) of the four feedinggroups were determined at days 14 and 49. Atday 14, two chambers in each group were ran-domly chosen and all fish in them were killed. Atday 49, all remaining fish were killed by overdosing

with anesthetic after their weights were measured.

Compensatory growth in black rockfish FISHERIES SCIENCE 847

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The fish were then freeze-dried and homogenized.Proximate analysis was conducted based on

AOAC14 methods. Protein [Kjeldahl method (N 6.25) after acid digestion], lipid (ether-extractionmethod), ash (muffle oven at 600C for 3 h) andmoisture contents (dry oven at 105C for 24 h) were

determined for the whole body samples. Energycontent was determined by a bomb calorimeter(PARR 1351, Moline, KS, USA).

Growth indices and statistical analysis

The following indices were calculated: specificgrowth rate (SGR, %/day) = 100 (lnWf- lnWi)/t,feeding rate (FR, % body weight/day) = 100 C/[(Wf+ Wi)/2]/t, and feed efficiency (FE, %) =100 (Wf- Wi)/C where Wf and Wi are final andinitial weights (g), t is the feeding duration (day)and Cis total feed consumption (g) duringtdays.

All statistical analyses were done using MinitabWindows v13 (Minitab, State College, PA, USA).Each replicate was considered as an experimentalunit, and thus, the mean value obtained from areplicate within each group was used as a data unit.Before day 14, all groups had five replicates (n = 5),and after then, three replicates were used in eachtreatment (n = 3). One-way analysis of variance(anova) was used to test the difference in the initialbody weight and chemical attributes between thefour groups, and then Tukeys multiple compari-son test was used when a difference occurred. Priorto anova, Levenes test was used to check homo-geneity of variance, and percents were arcsine-transformed. Analysis of covariance (ancova) withone factor was also used to compare the weeklymeasured body weight, SGR, FR and FE of the fourgroups. In ancova of body weight, the initial body

weight was used as a covariate to avoid anypossible confusion derived from the allometricrelationship between growth and body weight,although there was no difference in the initial body

weight between the four groups. In ancovaof SGR,FR and FE, body weight at the start of each week

was used as a covariate to compare the intrinsicability (i.e. growth potential) to return to the origi-nal growth trajectory. If needed, repeated measuresancovawas also done to analyze changes in growthand consumption over time using feeding treat-ments as between-subject, and days as within-subject. After ancova, planned contrasts amongadjusted means were done using an F-test (F= t2).

RESULTS

The body weight of juvenile black rockfish in thefour feeding treatments increased with time,

showing 100% survival rate during the wholeexperiment, but the weight obtained from the49-day cultivation was apparently different (Fig. 1).

At the beginning of the experiment, the weight ofthe fish undergoing the four treatments was notsignificantly different with a mean value of 1.43 g

(anova, P> 0.05). At day 14, however, there was asignificant difference in body weight betweenthe four treatments because of feed deprivation(ancova, P< 0.01); the mean weight of the control

was 3.43 g, but weights of F5, F10 and F14 were72.3, 51.0 and 35.2% of the control. Furthermore,this difference was continuously observed by theend of day 35. At day 42, however, F5 dramaticallybegan to catch up the control, and there was nosignificant difference between them thereafter(ancova, P> 0.05). In contrast, F10 and F14 showedsignificantly lower values than the control and F5(ancova, P< 0.05).

The temporal change in SGR based on weeklymeasured body weights after refeeding was appar-ently different between the four treatments (Fig. 2).SGRs of the control and F5 continuously decreased

with time, whereas SGRs of F10 and F14 showed aunimodal sequence, in which their SGRs increasedduring the first 14 or 21 days after refeeding (i.e.days 28 or 35) and then decreased. In addition,ancovaon SGRs adjusted for the weight at the startof each week showed that the degree of SGR,meaning the intrinsic ability to return to theoriginal growth trajectory, was also significantly

Time (day)

Bodywe

ight(g)

0

2

4

6

8

10Control

F5

F10

F14

0 14 21 28 35 42 49

a

b

cd

a

a

a

a

a a

b

b

b b

b

c

c

cc

c

d

d

a

d

a a a a

Fig. 1 Changes in body weight of black rockfish sub-jected to four feeding treatments during the 49-dayfeeding trial. Control, fish fed continuously; F5, fishfasted for 5 days (days 1014) and then fed to satiation;F10, fish fasted for 10 days (days 514) and then fed tosatiation; F14, fish fasted for 14 days (days 114) andthen fed to satiation. Values with different letters in thesame day are significantly different (P< 0.05). Data aremean standard error.

848 FISHERIES SCIENCE S-Y Oh et al.



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different between the treatments, and there was atendency that SGR increased with increments ofthe duration of feed deprivation except day 21. Atday 21, F5 showed significantly higher SGR thanthe other three groups (P< 0.05), and there was nosignificant difference between the three groups(P> 0.05). At day 28, however, SGRs of F5, F10 andF14 were significantly higher than that of thecontrol (P< 0.05), and both F10 and F14 showed

significantly higher SGRs than F5 (P< 0.05). Par-ticularly, the significantly higher SGRs of F10 andF14 compared to that of the control were continu-ously observed by the end of the experiment, butthe significantly higher SGRs of F5 compared tothat of the control were found only at days 28 and42 during the period between days 28 and 49.These results indicate that all black rockfish juve-niles fasted for 514 days exhibit compensatorygrowth after refeeding, but timing and degree ofcompensatory growth vary depending on the dura-tion of feed deprivation. Furthermore, whenFigures 1 and 2 are taken together, it is suggested

that complete compensatory growth can beachieved only in fish fasted for 5 days, whereas fishfasted for more than 10 days exhibit only partialcompensatory growth.

Although the temporal change in FR of F14 wasslightly different from that of SGR (Spearmansrank correlation, rs = 0.7), the correlation coeffi-cients (rs) between SGR and FR of the other threegroups were greater than 0.9, suggesting that SGRmay vary depending on FR (Fig. 3). Like SGR, FRalso showed a tendency to increase with the incre-ment of the duration of feed deprivation except

day 21, on which there was no significant differ-ence in FR between the four treatments.

FEs of the four feeding treatments during thewhole refeeding period ranged 55.899.7%, butthere was no significant difference between them(repeated measuresancova, between subject, feed-ing treatments P> 0.05; within subject, days P>0.05), suggesting that FE is not the factor affectingSGR (Fig. 4).anova for the chemical attributes (protein, lipid,

energy, ash and moisture content, and lipid/LBM)measured at day 14 showed that there were

Time (day)

21 28 35 42 49

Specificgrowthrate(%p

erday)

0

1

2

3

4

5

6

7

a

b

ab

a

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b

cc

a

abbc

c

a

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b

Control F5 F10 F14

Fig. 2 Changes in specific growth rate (SGR) of blackrockfish in four feeding groups during refeeding fromdays 1549. Feeding treatment abbreviations shown inFig. 1. Values with different letters in the same day are

significantly different (P< 0.05). Data are meanstandard error.

Time (day)

21 28 35 42 49

Feedingrate

(%p

erday)

0

1

2

3

4

5

6

7

a

b

c

d

a

abb

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aab

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Control F5 F10 F14

a

a aa

Fig. 3 Changes in feeding rate (FR) of black rockfish infour feeding groups during refeeding from days 1549.Feeding treatment abbreviations shown in Fig. 1. Valueswith different letters in the same day are significantly

different (P< 0.05). Data are mean standard error.

Time (day)

21 28 35 42 49

F

eedefficiency(%)

0

20

40

60

80

100

a

aa

a

aa

aa

a

a aa

a

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a a a

Control F5 F10 F14

Fig. 4 Changes in feed efficiency (FE) of black rockfishin four feeding groups during refeeding from days 1549.Feeding treatment abbreviations shown in Fig. 1. Valueswith different letters in the same day are significantlydifferent (P< 0.05). Data are mean standard error.




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significant differences between the four feedingtreatments (Table 1). Protein, lipid, and energycontent and lipid/LBM of F10 and F14 were signifi-cantly lower than those of control and F5 (exceptfor protein of F5 and F10, P< 0.01), but there wasno significant difference between the control andF5 in protein, lipid and lipid/LBM (P> 0.05), andonly the energy content of F5 was significantlylower than that of the control (P< 0.05). Likewise,ash and moisture content of F10 and F14 were sig-

nificantly higher than those of control and F5(P< 0.05), but there was no significant differencebetween the control and F5 in moisture content(P> 0.05). At day 49, however, there was no signifi-cant difference in energy, ash and moisture contentbetween the four feeding groups (P< 0.05). Fur-thermore, F10, which exhibited significantly lowervalues in protein content and lipid/LBM comparedto the control at day 14, dramatically caught up thecontrol. Similarly, F14 also caught up the control inprotein content, but its lipid/LBM was significantlylower than the control (P< 0.05).

DISCUSSION

Compensatory growth of fish after feed deprivationhas many advantages in aquaculture, includingefficient feed utilization and/or enhanced growthrate, minimized food waste and more flexiblefeeding regimes,15,16 but there have been a numberof inconsistent results. Some fish (e.g. channel cat-fish6 and Atlantic salmon7) have been reported toexhibit complete compensation, while others (e.g.

Alaska yellowfin sole8 and hybrid tilapia9) show

partial compensation. Furthermore, in certain fish(e.g. juvenile hybrid sunfish3), overcompensationhas been reported. In this study, however, com-plete and partial compensation co-occurred in thesame species, and the compensation pattern

was highly dependent on the duration of feeddeprivation.

Complete compensation is the most commonphenomenon in fish (e.g. rainbow trout Oncor-hynchus mykiss,2,17 Chinese longsnout catfish

Leiocassis longirostris,4 channel catfish Ictaluruspunctatus,6 minnow Phoxinus phoxinus,12 Atlanticcod Gadus morhua,13 hybrid tilapia Oreochromismossambicus O. niloticus,18 and Japanese floun-der Paralichthys olivaceus19). Furthermore, in thesespecies, fish fasted for more than 2 weeks areknown to exhibit complete compensation. Unlikethese species, however, in the juvenile black rock-fish used in this study, complete compensatorygrowth was achieved only in fish that experienced5-day feed deprivation (F5), but fish subjected tofeed deprivation for more than 10 days (F10 andF14) exhibited partial compensation. There might

be several factors to cause this difference betweenF5, F10 and F14. One possible factor was the rela-tive body mass of fasted fish to that of the controlfish. Tian and Qin15 reported that the weight of bar-ramundi Lates calcariferafter feed deprivation for1, 2 and 3 weeks was 63, 32 and 14% of the controland complete compensatory growth was onlyachieved in fish subjected to 1-week feed depriva-tion, thereby suggesting that the fasted fishthat experienced body mass below 60% of thecontrol may not exhibit complete compensation.Likewise, several studies have also reported such

Table 1 Body composition and energy concentration of black rockfish at day 14 and day 49 in four feeding groups

Date Chemical attribute

Treatments

C F5 F10 F14

Day 14 (n = 2) Protein 14.98 0.05a 14.95 0.14ab 14.24 0.05b 13.42 0.20c

Lipid 4.81 0.13a 4.49 0.03a 3.48 0.09b 2.19 0.09c

Energy 5.74 0.04a

4.85 0.01b

4.39 0.02c

4.34 0.01c

Lipid/LBM 0.25 0.01a 0.22 0.01a 0.18 0.01b 0.11 0.01c

Ash 4.33 0.02a 5.08 0.01b 5.41 0.02c 5.84 0.10d

Moisture 74.95 0.09a 75.21 0.23a 77.06 0.23b 78.37 0.44b

Day 49 (n = 3) Protein 14.98 0.09ab 14.69 0.09a 14.84 0.08a 15.53 0.15b

Lipid 7.30 0.04a 7.17 0.05ab 6.86 0.11bc 6.57 0.06c

Energy 6.59 0.05a 6.87 0.13a 6.61 0.07a 6.84 0.15a

Lipid/LBM 0.38 0.01a 0.38 0.01a 0.36 0.01a 0.33 0.01b

Ash 4.18 0.06a 4.11 0.14a 4.16 0.05a 4.17 0.08a

Moisture 72.30 0.17a 72.99 0.12a 72.92 0.51a 72.99 0.48a

Lipid/LBM, ratio of lipid to sum of protein and ash.C, control, fish fed continuously; F5, fish fasted for 5 days (days 1014) and then fed to satiation; F10, fish fasted for 10 days

(days 514) and then fed to satiation; F14, fish fasted for 14 days (days 114) and then fed to satiation.Values with different letters in the same row are significantly different (P< 0.05). Data are mean standard error.

Body composition (%, wet weight basis), energy concentration (kJ/g).




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weight-specific compensation growth. For ex-ample, Wang et al.18 reported that the weight ofhybrid tilapia after feed deprivation for 1, 2 and4 weeks was 71, 48 and 26% of the control, andcomplete compensatory growth was achieved onlyin fish subjected to 1-week feed deprivation. In

addition, Xie et al.20

reported that the weight ofgibel carp Carassius auratus gibelio after feed dep-rivation for 1 and 2 weeks was 92 and 74% ofthe control fish, and thus, complete compensatorygrowth was achieved in all groups within 2 weeksafter refeeding. Taken together, the result that com-plete compensatory growth was achieved only inF5 and there was no complete compensation inF10 and F14, which may be related to the relativebody mass of F5, F10 and F14 to that of the control(72, 51 and 35% for F5, F10 and F14, respectively).

Another possible factor was the physiologicalactivity to return to the original body mass after

refeeding. For example, although the basic energymetabolism decreased in all fasted groups, thedigestive organs and assimilation functions werethe most actively maintained in F5. This resulted inthe best feeding efficiency and growth perfor-mance within days after refeeding. In contrast,the feed-deprivation period of more than 10 daysmight cause serious adverse effects on digestiveorgans and their assimilation functions, therebyresulting in only partial compensation of F10and F14.

In this study, SGR of the black rockfish juvenilesfasted for 5 days (F5) was the highest at day 21

(7 days after refeeding),in which it was significantlyhigher than the control. Similarly, Zhu et al.21

reported that compensation growth in sticklebackGasterosteusaculeatusoccurredwithin 1 week afterrefeeding. In addition, the significantly higher SGRof F5 compared to the control lasted until day 28,disappeared at day 35, reappeared at day 42 andthen disappeared at day 49. This intermittentpattern in SGR is also shown in Tian and Qin,15 in

which 1-week fasted barramundi before refeedingfor 5 weeks exhibited an intermittent pattern inSGR. As pointed out inTian and Qin,15 this intermit-tent pattern in SGR is thought to be related to the

reduction of appetite after overfeeding in the previ-ous week, but further research is needed to eluci-date the relationship between them.

Inagreement withseveral studies,9,15we observedthat the significantlyhigher SGRs of F5, F10 andF14than that of the control decreased with time aftershowing a maximum value. Ingeneral,the time thatcompensation growth lasts after refeeding variesdepending on the experimental species. Forexample, compensatory growth lasts less than3 weeks in theminnow12 and two cyprinid species,22

but 8 weeks in juvenile Arctic charr Salvelinus alpi-

nus.11 In Atlantic salmon Salmo salar, compensa-tory growth lasts 80215 days after refeeding.7

Hyperphagia are considered to be a mainmechanism leading to compensatory growth offish during the refeeding period,3,4,6,16,18,23,24 whileimproved FE has not always reported in fish

achieving compensatory growth.2,4,12,25,26

In thisstudy, hyperphagia was observed in all fastedgroups after refeeding, but there was no significantdifference in FE among all feeding treatmentsduring the whole refeeding period. Therefore,increased FR of juvenile black rockfish during therefeeding period played a major role in compensa-tory growth in this study.

Jobling and Johansen27 proposed the lipostaticmodel hypothesis using lipid/LBM as an indicatorto predict the timing of compensatory growth. Inthis model, the decrease in lipid/LBM results ina compensatory growth response, whereas the

recovery of lipid/LBM to the control level mayresult in the termination of compensatory growth.Johansen et al.28 also reported some evidence sup-porting this model in Atlantic salmon. However,results of compensatory growth in gibel carp,20 bar-ramundi15,16 and Chinese longsnout catfish4 did notsupport the lipostatic model. In this study, lipid/LBMs of F10 and F14 at day 14 were significantlylower than that of control, whereas there was nosignificant difference between the control and F5.Despite this, F5 showed significantly higher SGRthan the control (i.e. compensation growth) duringthe next week (i.e. day 21), whereas there was no

compensation growth in F10 and F14 during thisperiod. This result suggests that juvenile blackrockfish may be an exception to the lipostaticmodel. However, since we did not measure lipid/LBM weekly, further studies are needed to eluci-date the relationship between compensationgrowth and lipid/LBM of black rockfish juveniles.

Juvenile black rockfish that only experiencedfeed deprivation for 5 days achieve complete com-pensatory growth during the following 5-weekrefeeding, whereas fish fasted for more than10 days exhibit only partial compensatory growth.This compensation growth pattern is therefore

thought to be negligible compared to a normalfeeding strategy. This finding, however, can providean insight into a practical feeding regime for blackrockfish farmers who want to fast fish for a certainperiod in order to save labor costs and for othereconomic reasons.

ACKNOWLEDGMENTS

Thanks to D-M Choi for experimental assistance.Funding was provided by the Korea Ocean




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Research and Development Institute throughproject PE98240.

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