JOURNAL OF THE Vol. 29, No.

3
WORLD AQUACULTURE SOCIETY September, 1998

Protein and Energy Levels for Maximum Growth, Diet

Utilization, Yield of Edible Flesh and Protein Sparing of
Hybrid Clarias Catfish {Clarias macrocephalus u
Clarias gariepinuc)
WIMOL JANTRAROTAI', PRASERT SITASIT, PATTANEE JANTRAROTA1,
THAV EE VIPUTHANUMA S AND KNEE SRABUA
National Inland Fisheries Institute, Bangkok 10900 Thailand

Abstract
Two feeding trials were performed to determine dietary protein and energy levels for
maximum growth, diet utillzation, yield of edible flesh and protein sparing of hybrid Clerics
*> ( <‘** *^***R !** ^ rias ganepinus). A 3 x 3 factorial deslgn was employed
for experiment 1 in which nine diets containing either 20, 30 or 409c protein and 275, 300
or 325-heel digestible energyfl00 g were used. The follow-up experiment 2 was a 3 x 2
factorial in which three protein levels (30, 35 and 40 to) and two digestible energy levels (275
and 325 kcal/100 g) were tested. Dletary ingredlents used in both studies were similar but
the ratios of energy provided by dextrin and lard were 11.3/1.0 and 3.8f1.0 for experiments
1 and 2, respectively. Each of the experimental diets was fed to apparent satiation twice daily
to three aquaria of fish (15 flsh/aquaria) with average weight of 4.0 and 2.0 g for experiment
1 and 2, respectively, for 8 wk. Performance of fish in experiment 1 was mainly influenced
by dietary protein rather than by energy levels. Maximum growth and diet utilization were
attained at dletary protein and energy levels of 409c and 275 kcal/100 g, respectively. Energy
levels did not spare dietary protein in experiment 1. In experiment 2, growth, diet utilization,
and yield of edible flesh of fish were influenced by protein and energy levels. Furthermore,
the protein sparing effect was observed in experiment 2. Best performance and protein spar-
ing were found at 359c protein and 325-kcal dlgestlble energyfl00 g. The studies showed that
dietary proteln and energy requirement for hybrid Clarias catfish are affected by carbohy-
drate to lipid ratio. Fish utilize dietary energy with a dextrin to lord ratlo of 3.8/1.0 more
emciently than that wlth a ratio of 11.3/1.0. Therefore, dietary protein and energy for best
performance and protein sparing of hybrid Clarias catfish reported for experiment 2 were
more appropriate.

Hybrid Clarias catfish (Clarias macro-
cephalus X Clarias gariepinus) are pres-
ently the most intensively cultivated fresh-
water fish in Thailand. Diets for the fish are
mostly extruded, contain 25 to 30% protein,
and are based on the requirements for na-
tive walking catfish C. batrachus (Chua-
poehuk 1987). In practice, hybrid Clarias
catfish fed these diets reach a marketable
size (100 g or more) in 90 to 120 d. How-
ever, hybrid Clarias catfish raised under
laboratory conditions can grow up to 100 g
in 70 d when fed with a diet containing
409c protein, 2.8 kcal/g digestible energy
(DE) (Jantrarotai et al. 1996). This may in-
' Corresponding author.
& Copyright by the Wotld Aquacuhure Society 1998
dicate that protein requirements for hybrid
Clarias catfish are more similar to that of
their male parent, C. gariepinus, rather than
that of native walking catfish (Degani et al.
1989).
Although 40% protein diets produce op-
timum growth of hybrid Clarias catfish, the
diets may not be cost effective as hybrid
Clarias catfish has low marketable value
(0.8—1.0 US$fkg). For other species, the op-
timum protein level of the diet can be low-
ered if the energy level is increased. due to
protein-sparing action of energy nutrients
(Lee and Putnam 1973; Machiels and Hen-
ken 1985; Serrano et al. 1992; Shiau and
Peng 1993). However, increasing energy in
fish diets may decrease feed efficiency and

281
282 I ANTRAROTAI ET Al.

TABLE 1. Comf›osition of the experimental diets (W• dry matter) used in experiment 1.

20% Protein 309• Protein 409• Protein

Energy (kcal/100 g) Energy (kcaf 100 g) Energy (kcal/100 g)
Composition 275 300 325 275 300 325 275 300 325
Ingredient
Peruvian fish meal 27.0 27.0 27.0 40.6 40.6 40.6 54.1 54.1 54.1
Fish oil 2.1 2.1 2.1 1.2 1.2 1.2 0.2 0.2 0.2
Soybean oil 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0
Dextrin 37.6 47.7 57.0 27.4 36.5 46.1 16.6 25.9 35.2
Lard 3.3 4.2 5.0 2.4 3.2 4.1 I.5 2.3 3.1
Vitamin & mineral mix' 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0
Vitamin C o. i 0.1 0.1 o. i o. i o.1 o. i o. i o. i
Cellulose 22.9 1 1.9 1.8 21.3 11 .4 0.9 20.5 10.4 0.3
Proximate composition
Protein 21.7 21.5 21.5 30.5 31.1 31.6 39.8 40.5 40.9
Lipid 10.7 11.4 11.7 9.9 10.5 11.2 8.7 9.5 10.2
Fiber 17.4 9.9 2.7 17.3 9.9 1.6 17.0 9.8 9.8
Ash 5.4 5.3 5.3 7.3 7.6 7.6 9.9 9.8 9.8
NFE 44.8 51.9 58.8 35.0 40.9 48.0 24.6 30.4 37.6
Digestible energy
(kcaU100 g) 275 297 324 275 296 322 271 295 320
Dextrin/lard 11.3 11.3 11.3 11.3 11.3 I 1.3 I 1.3 11.3 11.3
' Vitamin mixture provided the following per kg diet: vitamin A, 4,000 IU; D„ 2,000 IU; vitamin E, 50 IU;
menadione sodium bisulfite, 10 mg; thiamin, 20 mg; riboflavin, 20 mg; niacin, 150 mg; calcium pantothenate,
200 mg; folic acid, 5 mg; pyridoxine, 20 mg; choline chloride. 2,000 mg; vitamin B, z, 0.2 mg; biotin, 2 mg;
and inositol, 400 mg. Mineral mixture provided the following per kg diet: manganese, 25 mg; zinc, 20 mg;
copper. 5 mg; iodine, 5 mg; cobalt 0.05 mg; selenium, 0.3 mg; and iron, 30 mg.

lead to increased fat deposition (Reis et al.
1989). Protein-to-energy ratios in fish diets
must be carefully manipulated. The purpose
of this study is to determine different di-
etary protein and energy levels on growth,
diet utilization, yield of edible flesh and
protein sparing of hybrid Clarias catfish.
Materials and Methods
Experimental Conditions
Hybrid Clarias catfish fingerlings were
obtained from a commercial hatchery and
acclimated to laboratory conditions for 2
wk in a 2,000-L rectangular tank with con-
tinuously flowing water (1 Lfmin). During
the acclimation period, they were fed a
sinking-type feed pellet prepared from com-
mercial catfish feed (309c protein). Two ex-
periments were designed to study the dif-
ferent dietary protein and energy levels on
performance and protein sparing of the fish.
Twenty-seven 46W X 91L X 46H cm and
18 35W X 60L X 35H cm glass aquaria
were used in experiments 1 and 2, respec-
tively. The aquaria were supplied with aer-
ation and overflow pipes that allowed water
to flow through at the rate of 0.8 L/min.
Well water was supplied from an outdoor
tank by gravity flow to the laboratory. Wa-
ter temperature was maintained at 28 1 2
C throughout the experiments. Fluorescent
lights illuminated the laboratory for 14 h
per d, 0600 to 2000 h.
Experimental Design and Diets
A 3 X 3 factorial design was employed
for experiment 1 in which nine diets were
formulated to contain either 20, 30 or 409a
protein and 275, 300 or 325 kcal digestible
energy (DE)f1O0 g diet (Table 1). The diets
contained Peruvian fish meal as sole protein
source, fish oil and soybean oil to provide
essential fatty acids, and a mixture of dex-
trin and lard at weight ratio of 11.3/1.0 to
 PROTEIN AND ENERGY lN CURIAS DIETS 283

TABLE 2. Comf›osition r›/ the experimental diets (to dry matter} used in experiment 2.

309c Protein 359c Protein 409• Protein

Energy (kcal/100 g) Energy (kcal/100 g) Energy (kcal/100 g)
Composition 275 325 275 325 275 325
Ingredient
Peruvian fish meal 40.6 40.6 47.3 47.3 54.1 54. I
Fish oil 1.2 1.2 0.7 0.7 0.2 0.2
Soybean oil 3.0 3.0 3.0 3.0 3.0 3.0
Dextrin 18.8 33.0 15.6 29.7 12.4 26.6
Lard 4.9 8.6 4.1 7.8 3.2 7.0
Vitamin & mineral mix' 4.0 4.0 4.0 4.0 4.0 4.0
Vitamin C 0.1 0.1 0.1 0.1 0.1 0. i
Cellulose 27.4 9.5 25.2 7.4 23.0 5.0
Proximate composition
Protein 30.8 30.3 35.7 35.3 40.9 40.9
Lipid 12.1 15.7 11.6 15.0 10.4 13.9
Fiber 24.8 10.7 21.1 8.8 21.1 9.6
Ash 7.3 7.4 8.5 8.7 9.7 9.6
NFE 25.0 35.9 23.1 32.2 17.9 26.0
Digestible energy
(kcal/100 g) 268 323 276 325.5 273 321
Dextrin/lard 3.8 3.8 3.8 3.8 3.8 3.8
' Vitamin mixture provided the following per kg diet: vitamin A, 4,000 IU; D„ 2,000 IU; vitamin E, 50 IU;
menadione sodium bisulfite, 10 mg; thiamin, 20 mg; riboflavin, 20 mg; niacin, 150 mg; calcium pantothenate,
200 mg; folic acid, 5 mg; pyridoxine, 20 mg; choline chloride. 2.000 mg; vitamin , 0.2 mg; biotin, 2 mg;
and inositol. 400 mg. Mineral mixture provided the following per kg diet: manganese, 25 mg; zinc, 20 mg;
copper, 5 mg; iodine. 5 mg; cobalt 0.05 mg; selenium. 0.3 mg; and iron, 30 mg.

balance required DE contents. The follow-
up experiment 2 was a 3 X 2 factorial in
which three protein levels (30, 35 and 409c)
and two DE levels (275 and 325 kcaf 100
g) were tested. The diets in experiment 2
(Table 2) were similar to those used in ex-
periment l except they contained dextrin
and lard at a weight ratio of 3.8/1.0. The
DE of the diets was estimated using
lished DE values for channel catfish, 3.5
kcal/g protein, 8.1 kcal/g lipid and 2.5 kcal/
g carbohydrate (NRC 1993). The diets were
prepared and analyzed with the same pro-
cedures described by Jantrarotai et al.
(1994).
Feeding Trial
Each of the experimental diets was ran-
domly assigned to three aquaria of hybrid
Clarias catfish. Each aquarium contained
15 fish with an average initial weight of 4.0
* 0.04 and 2.0 * 0.00 g/fish for experi-
ments 1 and 2, respectively. Fish were fed
the experimental diets in each aquarium to
apparent satiation twice daily, at 0800 and
1700 h for 8 wk. Uneaten diet in each
aquarium was siphoned and collected for
determination of dried weight after 1 h of
feeding. Fish were weighed and counted bi-
weekly.
pub-
Sample Collection and Analyses
At the beginning of the experiment, a
representative sample of fish was collected
for initial whole body composition analysis.
At the end of the experiment, 10 fish from
each aquarium were randomly collected.
Five fish were used for final whole body
composition analysis. The analyses were
conducted according to standard methods
(AOAC 1984). Energy of whole fish was
calculated using gross energy value, 5.65
and 9.45 kcafg protein and lipid, respec-
tively. Another five fish were sacrificed and
284 JANTRAROTAI ET AI.

T BkE 3. Growth performance and diet utilization of hybrid Clarias cotfish fed diets with di erent f›rotein and
energ y levels in experiment I. Values are means from three groups of fish. Italic numbers are the average
values of main factors from nine groups of fish. Different letters abc and xy indicate significant (P < 0.05)
difference.s within protein and energ y effects, respectively.

209o Protein 309c Protein

Energy (kcal/100 g) Energy (kcal/100 g)
Variable 275 300 325 275 300 325
Mean initial weight (g) 4.10 4.00 4.10 4.10 4.10 4.10
Mean final weight (g) 48.10 50.80 69.10 79.40 84.40 86.50
Weight gain (to) 1,009 1,170 1,625 1,883 2,008 2,060
Feed efficiency ratio 0.41 0.38 0.38 0.58 0.52 0.56
Protein efficiency ratio 1.89 1.79 1.82 2.02 1.70 1.79
Apparent protein retention ('to) 31.80 29.40 29.50 32.20 29.50 30.80
Utilization of digestible energy (to) 29.60 29.00 31.30 42.00 39.00 36.50
Feed intake (to BW/d) 7.42 7.96 7.29 5.54 6.23 5.37
Protein intake (to BW/d) 1.61 1.71 1.56 1.69 1.94 1.70
Energy intake (kcal/100 g per d) 20.33 23.65 23.62 15.19 18.44 17.25

excised for liver and whole muscle for the Protein efficiency ratio = weight gain/pro-
determination of hepatosomatic index (HSI) tein intake
and yield of edible flesh. The remaining Apparent protein retention (%) = [(final
pan of the fish after liver and muscle were body protein — initial body protein)/
excised was weighed for the determination protein intake] X 100
of percentage of carcass waste. Utilization of digestible energy (%) = [(fi-
nal body energy — initial body ener-
Data Calculation and Analysis gy)/energy intake] X 100
The following parameters were calculat- Feed intake (to BW/d) = (mean daily feed
ed for evaluation of hsh performance: intake per fish N 100)/[(initial weight
+ final weight)/2]
Weight gain (g/fish) = final weight — initial Protein intake (to BW/d) = (feed intake) X
weight (dietary protein/g)
Feed efficiency ratio = weight gain/feed in- Energy intake (kcafl00 g fish per d) =
take (feed intake) X (dietary energy/g)

TABLE 4. Biological parameters of hybrid Clarias catfish fed diets with different protein and energ y levels in
exf›eriment I. Values are means frum three groups of fish. italic numbers are the average vnJtie.r of main
factors from nine gr •R* • f fish. Di erent letter abc and xy indicate significant (P < 0.05) differences within
protein and energ y effects, respectively.

209c Protein 309• Protein

Energy (kcal/100 g) Energy (kcal/100 g)
Variable 275 300 325 275 300 325
Hepatosomatic index ('to) 1.53 1.24 1.39 t.2t 1.10 1.15
Flesh (to) 37.00 3730 38.80 38.20 38.70 38.60
Carcass waste (to) 56.90 5610 54.70 5710 55.10 56.50
Whole body composition (9•)
Moisture 67.30 66.80 64.60 68.60 67.60 67.50
Protein 50.90 49.00 49.50 53.30 53.20 53.00
Lipid 31.70 33.50 37.40 31.20 36.80 34.50
Ash 11.20 i i.vo io.so 12.70 12.10 11.30
 PROTEIN AND ENERGY IN CURIAS DIETS 285

TABLE 3. Extended.

409c Protein
Protein mean Energy mean
Energy (kcal/100 ,›
275 3D0 325 20 30 40 275 300 325
4.00 4.10 4.10 4. IOa d.lOa 4. IOa 4. lOx 4. 10x 4. IOx
109.00 108.40 102.90 56.OOc 83.dOb 106.80a 78.9Ox 81.2Ox 86.20a
2,625 2,608 2.470 I,298c I,983b 2,568a ?,869z I,929x 2,O52x
0.70 0.67 0.65 O.39c 0.55b 0.67a fi.56a 0.52a 0.53x
1.75 1.65 1.62 1.83a 1.84a 1.67a I.89x I.7Ix 1.74x
31.90 29.10 26.30 30.2Oa 30.9Oa 29. IOa 31.9Ox 29.3Ox 28.9Hz
49.20 41.80 36.80 29.90b 39.20a 42.60a dO.2Ox 36.60x 3d.9Ox
4.77 4.98 5.07 7.56a 5.71b 4.94b 5.91x 6.39x 5.9/
1.90 2.02 2.07 1.63b 1.77b 2.OOa 1.73x 1.89x I.78x
12.93 14.68 16.22 22.53a J6.966 l4.61c 16. lS y 18.92x 19.03x

Hepatosomatic index (') = (liver weight/
body weight) X 100
Edible flesh (9r) = (muscle weight/body
weight) X 100
Carcass waste (9«) = (fish waste weight/
body weight) X 100
Data were analyzed by factorial analysis
of variance to determine the main effects of
protein and energy level and the effects of
interactions between the two factors. Ho-
mogeneity of variance was confined using
Cochran’s Q test. Data which expressed as
percentage or ratio were subjected to arc-
sine transformation prior to analysis. When
a significant main effect was found (I <
0.05), mean differences were determined by
Duncan’s multiple range test. Statistical
analysis was performed using Statgraphics
software.
Results
No interaction (P > 0.05) between pro-
tein and energy levels was found for any of
the variables determined in both experi-
ments. In experiment 1, increasing protein
in the diets from 20 to 409c improved
weight gain of fish and feed efficiency (I
< 0.05) (Table 3). Energy retention was
higher (P < 0.05) for fish fed 30 and 409»
protein than for fish fed 209c protein (Table
3). Protein efficiency ratio and protein re-
tention were not affected {P > 0.05) by di-
etary protein. Dietary energy levels had no
influence (f• > 0.05) on weight gain, feed

TABLE 4. Extended.

409c Protein
Energy (kcal/100 g) Protein mean Energy mean
275 300 325 20 30 40 275 30D 325
1.07 1.03 1.07 1.38a 1.16b I.05b 1.26a 1.1z i.zoo
40.20 41.80 40.00 37.70b 38.50b 40.90a 38.40x 39.3Ox 39.40x
54.70 53.10 54.50 55.90a 56.20a 54.101› 56.20x 54.80y 55.20xy

68.30 68.50 70.50 62.20c 67.90b 69.10a 68.I0x 67.60x 67.50x

57.10 55.60 52.20 48.60c 53.20b 56.His 53.80x 52.6Oxy 5I.d0y
28.80 27.80 30.00 34.20a 34.20a 28.90b 30.60x 32.70x 34.00x
12.00 11.60 11.90 I?.20n J2.Ibn 11.80a 11.90x 11.80x 11.30x
286 JANTRAROTAI ET Al.

TABLE 5. Growth performance and diet utilization of hybrid Clarias catfi.sh fed diets with different protein and
energ y levels in experiment 2. Values are means from three groups of fish. Italic numbers are the average
values of main factors from nine groups of fish. Different lelcers ahc and xy indicate significant (P < 0.OS}
differences within protein and energ y effects, respectively.

309c Protein 359c Protein 409c Protein

Energy (kcal/100 g) Energy (kcal/100 g) Energy (kcal/100 g)
Variable 275 325 275 325 275 325
Mean initial weight (g) 2.00 2.00 2.00 2.00 2.00 2.00
Mean final weight (g) 33.80 61.60 54.30 77.30 59.00 77.30
Weight gain (to) 1.589 2,978 2,014 3,738 2,841 3,767
Feed efficiency ratio 0.98 0.97 1.01 1.06 0.99 1.08
Protein efficiency ratio 3.17 3.21 2.84 3.05 2.42 2.63
Apparent protein retention (to) 55.00 57.20 51.90 52.20 39.40 44.90
Utilization of digestible energy (to) 69.70 64.30 73.50 66.70 66.50 66.40
Feed intake (9• BW/d) 3.20 3.40 3.30 3.20 3.36 3.15
Protein intake (9• BW/d) 0.98 1.04 1.17 1.12 1.38 1.29
Energy intake (kcal/100 g per d) 8.50 11.13 9.10 10.37 9.20 10.17

and protein efficiency ratio, and apparent In experiment 2, dietary protein and en-
protein and energy retention. Fish fed 30 ergy independently affected growth of fish
and 40% protein diets had lower daily feed (Table 5). Increasing dietary protein at each
intake (I- < 0.05) than those fed 209» pro- energy level from 309c to 35% improved
tein diets (Table 3). However, daily protein weight gain of fish (P < 0.05). Fish fed
intake of fish fed 20 and 309» protein was 409c protein diets did not grow faster (I' >
not different (I 0.5) but was less (P < 0.05) than those fed 35% protein diets with
0.05) than for fish fed 409c protein. Daily the same energy levels. Increasing dietary
energy intake was affected by both dietary energy at any protein level significantly (I
protein and energy levels (Table 3). Daily < 0.05) improved fish growth. Feed effi-
energy intake decreased (I- < 0.05) as di- ciency ratios were high in all dietary treat-
etary protein increased. Fish fed low energy ments and were not affected (I > 0.05) by
diets had less (P < 0.05) daily energy in- dietary protein and energy levels (Table 5).
take than those fed higher energy levels. Protein efficiency ratios and apparent pro-
Fish fed 209c protein diets had higher (I < tein retentions were high for diets contain-
0.05) HSI than fish fed higher protein levels ing 30 and 35% protein and decreased (I
(Table 4). Yield of edible fish flesh and car- < 0.05) for the higher protein diets. Energy
cass waste were inversely related. Fish fed levels had no effect (I > 0.05) on protein
409c protein diets had more edible flesh and efficiency ratio and apparent protein reten-
less carcass waste than the other groups (P tion (Table 5). Utilization of digestible en-
< 0.05). Carcass waste tended to increase ergy was not affected (P > 0.05) by dietary
for the fish fed diets containing energy lev- treatments. Daily feed intake of fish in ex-
els of 275 kcal/100 g. Moismre, protein and periment 2 was similar (P > 0.05) for all
lipid in whole fish were affected by dietary treatments (Table 5). However, protein and
protein but only protein content was affect- energy intake increased as dietary protein
ed by dietary energy. Increasing dietary and energy increased, respectively. Fish fed
protein caused increased (P < 0.05) whole 309c protein diets had higher (P < 0.05)
body moisture and protein content but tend- HSI than fish fed higher protein levels. En-
ed to decrease lipid content (Table 4). Pro- ergy levels had no influence {P > 0.05) on
tein content of whole body also decreased HSI (Table 6). Yield of edible flesh, carcass
(P < 0.05) as dietary energy increased. waste and whole body composition (Table
 PROTEIN AND ENERGY IN CURIAS DIETS 287

TABLE 5. Extende‹J.

Protein mean Energy mean

30 35 40 275 325
2.OOo 2.ODo 2.00a 2.OOH
47.70b 65.50a 68.10a 49.OOy 71.90x
2,284b 3,176a 3,304a 2,3d8y 3,494x
0. 98a /.O4n /.04o 0. 99x /.04z
3.19a 2.94a 2.52b 2.8Ix 2. V6
56.10a 52. 10a 42.20b 48.8Ox 51.4Ox
67.00a 7O.10a 66.50a 66.90x 65.8Ox
3.32a 3.23a 3.26a 3.29x 3.26z
1.Olc J. l5h 1.34a 1. l8x 1. I5x
9.82u 9.73a 9.68a R.9Oy 10.55x

6) of all fish were similar and not affected
[P > 0.05) by dietary protein and energy
levels.
Discussion
The best protein and energy for maxi-
mum weight gain, diet utilization and yield
of edible flesh of hybrid Clarias catfish in
experiment 1 was 40' r and 275-kcal DE/
100 g. These values are similar to those re-
ported for C. gariepinus (Machiels and
Henken 1985; Degani et al. 1989). Fish per-
formance in the present study was influ-
enced by dietary protein levels rather than
by energy levels. Machiels and Henken
(1985) related fish growth and feed effi-
ciency as functions of dietary protein in-
take. They found that increasing protein in-
take of C. gariepinus fed diets with differ-
ent protein levels improved growth and
feed efficiency. Our study clearly showed
that fish fed the 409c diets consumed more
protein than those fed lower protein diets
(Table 3) and therefore grew and utilized
feed better. Energy retention, which was
low in fish fed the low protein diets, is in

TABLE 6. Biological parameters of hybriJ Clarias catfish fed diets with different f›rotein and enerpy levels in
experiment 2. Values are mean.c from three groups af fish. Italic numbers ure the average valuec nf main
facior.s frnm nine 8•••r* of fish. Different letter abc and xy indicate significant (P < 0.OS} differences within
protein and energ y effects, respectively.

.30& Protein 35a Protein 40% Protein

Protein mean Energy mean
Energy (Kcal/HD g) Energy (Kcal/I(D g) Energy (Kcal/100 g)
Variable 275 325 275 325 275 325 30 35 40 275 325
Hepatosomatic
index (9•) 1.77 1.78 1.43 1.56 1.51 1.40 I.78a I.50b 1.45b 1.57x 1.58x
Flesh (to) 34.30 35.40 35.40 36.20 36.40 36.80 34.80a 35.80a 36.60a 35.60x 36.lOx
Carcass waste
(9c) 57.10 56.20 56.90 55.20 56.40 55.90 56.60 56.OOa 56. IO‹i 56.8Ox 56.8Ox
Whole body composition (9•)
Moisture 68.50 66.80 68.40 66.00 70.10 68.40 d7.70a 67.20a 69.3Oa d9.OOH 67.1Ox
Protein 54.20 53.10 57.40 53.20 53.80 53.70 53.70a 55.30a 53.ROa 55.IOx 53.3Ox
Lipid 31.50 35.50 31.40 33.50 31.90 33.70 33.50a 32.5Oa 32.80a 3I.6Ox 34.20x
Ash 10.70 10.80 11.60 11.30 13.00 11.10 10.80a 11.5Oa 12.10a ll.8Ox 11.lOx
288 JANTRAROTAI ET Al.
agreement with that reported for C. garie-
pinus (Machiels and Henken 1985; Degani
et al. 1989). However, for C. gariepinus, the
protein retention was also low for fish fed
the low protein diets leading Degani et al.
(1989) to conclude that some protein was
used for energy. This is in contrast to our
study where protein retention of fish fed the
low protein diets was similar to that for fish
fed higher protein diets. This is probably
because fish fed the low protein diets con-
sumed enough non-protein energy so that
the energy intake of fish in this group was
the highest (Table 3).
Increasing energy levels over 275 kcaU
100 g had no beneficial effects on fish per-
formance. This agrees with results for C.
gariepinus (Degani et al. 1985) and hybrid
striped bass ‹Morone chrysops X M. saxa-
tilis) (Nematipour et al. 1992). However,
studies with C. gariepinus (Machiels and
Henken 1985), channel catfish Ictalurus
punctatus (Garling and Wilson 1976), nile
tilapia Oreochromis niloticus (El-Sayed and
Teshima 1992) and rainbow trout Oncor-
yhnchus mykiss (Lee and Putnam 1973) had
shown that increasing both protein and en-
ergy to the optimum levels resulted in better
growth of fish and improved feed efficien-
cy. The difference may relate to energy
source. In our study, carbohydrate was the
major energy source whereas the others
used more lipid for energy. This assumption
was tested in experiment 2 in which diets
with a low ratio of dextrin to lard (3.8/1.0)
were used. In this experiment, optimum
growth, diet utilization and yield of edible
flesh of hybrid Clarias catfish were attained
at 35% protein and 325 kcal/100 g. The
protein level was lower and the energy level
was higher than the values reported in ex-
periment 1. Furthermore, the improved
growth rates as well as protein sparing ef-
fect were observed in fish fed higher energy
diets in experiment 2. These findings did
not present in experiment l when a high
ratio of dextrin and lard (11.3/1.0) was used
as energy source. The difference in protein
and energy requirement for hybrid Clarias
catfish in both experiments was clearly due
to the different ratios of carbohydrate and
lipid provided as dextrin and lard. The find-
ings indicated that hybrid Clarias catfish
utilized energy with a low ratio of carbo-
hydrate to lipid more efficiently and this re-
sulted in a protein sparing effect. Similarly,
Degani et al. (1989) found that dietary lipid
for C. gariepinus was used mostly as an
energy source and spare protein. Deviations
in digestible energy of dextrin for hybrid
Clarias catfish from the “average” carbo-
hydrate DE value of 2.5 kcal/g (NRC 1993)
for channel catfish may have reduced actual
differences in DE between diets in experi-
ment 1 and rendered no effect of the energy
on performances of fish.
Feed intake of fish in experiment 2 was
not regulated by either dietary protein or
energy levels since fish in all groups con-
sumed a similar amount of feed (Table 5).
This resulted in a greater intake of protein
and energy for fish fed higher protein and
energy levels. In this experiment, protein
efficiency and apparent protein retention
decreased when protein was above 359c.
According to Hepher (1988), protein effi-
ciency ratio and protein retention are usu-
ally low at low levels of dietary protein be-
cause certain amino acids may limit protein
utilization, which is optimal, decreasing
thereafter due to the utilization of a portion
of the protein for maintenance and the over-
all energy level of the ration. However, we
found that energy levels in experiment 2
had no effect on protein utilization. It could
be, therefore, 409c protein is above require-
ment and excess amino acids are catabo-
lized and deaminated.
Hepatosomatic index of fish in both ex-
periments was highest (I < 0.05) for those
fed the lowest protein level. This is proba-
bly due to liver lipid accumulation as a re-
sult of the lowest dietary protein to energy
ratios among all diets (Nematipour et a1.
1992; Serrano et al. 1992; Jantrarotai et al.
1994).
Fish fed 20 and 309c protein diets in ex-
periment 1 yielded markedly lower edible
 PROTEI N AND ENERGY IN CURIAS DIETS
flesh and higher carcass waste than those
fed 409c protein. By contrast, in experiment
2, in which a low ratio of dextrin to lard
was used, yield of edible flesh and whole
body protein of fish in all groups did not
differ suggesting that lipid spared protein
resulting in a greater deposition of protein
in fish fed 30 and 359c protein diets.
The study suggested that dietary protein
and energy requirement for hybrid Clarias
catfish are affected by carbohydrate to lipid
ratio. The fish utilize dietary energy with a
dextrin to lard ratio of 3.8/1.0 more effi-
ciently than that with a ratio of 11.3/1.0 and
hence spare dietary protein. Therefore, the
dietary protein and energy level with a low
ratio of carbohydrate to lipid for maximum
growth, diet utilization, yield of edible flesh
and protein sparing of hybrid Clarias cat-
fish are 359c and 325 kcal/100 g, respec-
tively.
Acknowledgments
This work was supported by the Austra-
lian Centre for International Agriculture
Research (ACIAR). We thank G.L. Allan
for reviewing the manuscript.
Literature Cited
AOAC (Association of Official Analytical Chem-
ists). 1984. Official methods of analysis, 14th edi-
tion. AOAC, Arlington, Virginia, USA.
Chuapoehuk, W. 1987. Protein requirements of walk-
ing catfish Clarias hairachus (Linnaeus), fry.
Aquaculture 63:215—219.
Degani, D., Y. Ben-Zvi and D. Levannon. 1989. The
effect of different protein levels and temperature
on feed utilization. growth and body composition
of Clarias gariepinus (Burchell 1822). Aquacul-
ture 76:293—30l.
El-Sayed, A. M. and S. Teshima. 1992. Protein and
289
energy requirements of Nile tilapia, Oreochromis
niloticus, fry. Aquaculture 103:55—63.
darling, D. L., Jr. and R. P. Wilson. 1976. Optimum
dietary protein to energy ratio for channel catfish
fi ngerlings Ictalurus punctatus. Journal of Nutri-
tion 106:1368—1375.
Hepher, B. 1988. Nutrition of pond fishes. Cambridge
University Press, New York. USA.
Jantrarotai, W., P. Sitasit and S. Rajchapakdee.
1994. The optimum carbohydrate to lipid ratio in
hybrid Clarias catfish (Clarius macrocephalus X
C. garieyinu.s) diets containing raw broken rice.
Aquaculture 127:6l —68.
Jantrarotai, W., P. Sitasit and A. Sermwatanakul.
1996. Quantifying dietary protein level for maxi-
mum growth and diet utilization of hybrid Clarias
catfish (Clarias macrocephalus X C. gariepinus).
Journal of Applied Aquaculture 6:71—79.
Lee, D. J. and G. B. Putnam. 1973. Response of rain-
bow trout to varying protein/energy ratio in a test
diet. Journal of Nutrition 103:91ñ—922.
Machiels, M. A. M. and A. M. Henken. 1985.
Growth rate, feed utilization and energy metabo-
lism of the African catfish, Clarias gariepinus
(Burchell. 1822) as affected by dietary protein and
energy content. Aquaculture 44:271—278.
Nematipour, G. R., M. L. Brown and D. M. Gatlin
III. 1992. Effects of dietary:protein ratio on
growth characteristics and body composition of
hybrid striped bass, Mnrone chry.Sof›.s K ñf. .v‹zx-
atilis. Aquaculture 107:359—368.
NRC (National Research Council). 1993. Nutrient
requirements of fish. National Academy Press,
Washington, D.C., USA.
Reis, L. M., E. M. Reutebuch and R. T. Lovell.
1989. Protein-to-energy ratios in production diets
and growth, feed conversion and body composi-
tion of channel catfish, Ictalurus punctatus. Aqua-
culture 77:21—27.
Serrano, J. A., G. R. Nematipour and D. M. Gattin
III. 1992. Dietary protein requirement of red
drum {Sciaenops ocellatus) and relative use of di-
etary carbohydrate and lipid. Aquaculture 101:
283—291.
Shiau, S. and C. Peng. 1993. Protein-sparing effect
by carbohydrates in diets for tilapia, Oreochromis
niloticus S 0. aureus. Aquaculture 117:327-334.