HAYATI Journal of Biosciences January 2015

Available online at:

Vol. 22 No. 1, p 12-19 http://journal.ipb.ac.id/index.php/hayati
EISSN: 2086-4094 DOI: 10.4308/hjb.22.1.12

Growth Response and Feed Utilization of

Giant Gourami (Osphronemus goramy) Juvenile
Feeding Different Protein Levels of the Diets Supplemented
with Recombinant Growth Hormone
DARMAWAN SETIA BUDI1*, ALIMUDDIN2, MUHAMMAD AGUS SUPRAYUDI2

1
Study Program of Aquaculture Science, Department of Aquaculture, Faculty of Fisheries and Marine Science,
Bogor Agricultural University, Dramaga Campus, Bogor 16680, Indonesia
2
Department of Aquaculture, Faculty of Fisheries and Marine Science, Bogor Agricultural University,
Dramaga Campus, Bogor 16680, Indonesia

Received June 12, 2014/Accepted December 11, 2014

The purpose of this study was to examine the effect of dietary supplementation with recombinant growth hormone
(rGH) on the growth and dietary utility of juvenile giant gourami. The rGH was mixed with chicken egg yolk and
sprayed on to artificial feed with different protein levels (34, 28, and 21%; isoenergy). Each treatment group of gourami
was paired with a control group that received feed of the same protein level, but without rGH supplementation.
Juvenile of giant gourami (weight 15.83 + 0.13 g) were fed diets containing rGH, to apparent satiation, 2 times a
week. Fish were reared from less than 2 months old for 42 days in 100 L glass aquaria at an initial density of 10 fish
per aquarium. At the end of this period, the biomass and daily growth rate (SGR) of the fish were measured and the
feed conversion ratio calculated and compared. Our data showed that fish fed rGH-supplemented diets experienced
higher growth than fish in the control groups and showed that fish with higher protein diets experienced higher growth
than the groups with less protein diets. The group with the highest biomass gain, SGR, and lowest feed conversion
ratio (FCR) was the group fed a 34% protein diet supplemented with rGH. Furthermore, biomass gain, SGR, and
FCR in the rGH treatment group with a 28% protein diet matched the measurements of the non-rGH control group
receiving a 34% protein diet (P > 0.05). We conclude that giant juvenile gourami given feed supplemented with
recombinant growth hormone will experience increased growth and dietary utility compared with gourami given
the same feed without supplementation.

Key words: feed protein levels, growth, giant gourami, rGH

___________________________________________________________________________

INTRODUCTION specifically, prior research has shown that rGH

application via immersion method significantly
Giant gourami (Osphronemus goramy) is an increases the growth rate of giant gourami (Irmawati
important aquaculture commodity in Indonesia. et al. 2012). The application of rGH with 3 mg/kg
Giant gourami grows at a relatively slow rate, dose in gourami feed also resulted in higher growth
which can hinder efforts to increase giant gourami than control groups without rGH (Safir 2012). This
production to meet high market demand. However, study aims to examine the combined effect of rGH
if increasing the production of giant gourami is application and variation of dietary protein levels,
difficult, it may be possible to increase the growth which has not yet been explored.
of each fish. There are several known strategies for The rGH can increase metabolic capacity,
increasing fish growth rate. The use of recombinant especially lipid metabolism and optimization of
growth hormone (rGH) in fish diet is an easy protein synthesis, on giant gourami. Fish with a
method that can yield quick results in fish growth. faster relative growth rate exhibited the protein
Many studies have examined the effect of rGH sparring effect that occurs with increased growth
supplementation on fish species such as Nile tilapia hormone levels (Rasmussen et al. 2001; Irmawati et
(Acosta et al. 2007), rainbow trout (Llorente et al. al. 2012). Growth and development of body tissue
2004), abalone (Moriyama et al. 2008), and yellow in fish are influenced by the balance of protein and
tail (Pedroso et al. 2009). Regarding giant gourami
_________________
energy in their feed. Feed with high protein content

Corresponding author. Phone: +6285649871678
∗ will not necessarily accelerate growth if the total
Fax: +62-251-8622941, E-mail: darma.jhders@gmail.com energy content is low. The energy content of feed is
Copyright © 2015 Institut Pertanian Bogor. Production and hosting by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
Vol. 22, 2015 Growth of Giant Gourami Feeding with Recombinant Growth Hormone �������
13

mainly used for primary metabolic activities, such as The rGH was coated with 20 mg chicken egg yolk
respiration, ion transport/metabolites, regulation of and then applied to feed pellets at a dose of 3 mg/kg
body temperature and other physiological functions. (Safir 2012). The 3 treatment groups that received
Energy for these activities generally derives from feedat each protein level, coated with rGH, were
non-protein nutrients (fats and carbohydrates). If paired with control groups that received the feed at
the energy contribution from non-protein material the same protein level but without rGH, for a total of
is relatively low, protein must be tapped to produce 6 experimental groups. We conducted 3 repetitions
energy. This reduces the amount of protein of the experiment for all 6 groups.
available for building body tissues. In other words, Fish Rearing. Juvenile of giant gourami (BW
the addition of non-protein nutrients for energy 15.83 + 0.13 g) were reared for 42 days in 100 L glass
production can minimize the use of protein as an aquaria at initial density of 10 fish per aquarium.
energy source and maximize its use for growth and Water was changed 50% per day each afternoon.
body mass (protein sparring effect). This improves Oxygen levels were maintained using aeration. Fish
the function of protein in supporting the growth were fed three times a day (morning, afternoon, and
of fish (Furuichi 1988; Hasan & Khan 2013). The evening) until apparent satiation. Treatment groups
rGH supplementation is expected to increase the were given feed containing rGH 2 times a week, at
efficiency of protein utilization in the diet. The an interval of 3 days.
purpose of this study was to examine the effect of Data Collection and Statistical Analysis. Fish
different dietary protein levels supplemented with body weight was measured every 14 days until
recombinant growth hormone (rGH) on the growth at the end of the experiment. For each fish, we
and dietary utility of juvenile giant gourami. measured hepatosomatic index (HSI), proximate
composition, excreted total ammonia nitrogen
MATERIALS AND METHODS (TAN), blood glucose and triglycerides levels, liver
and muscle glycogen, and enzyme activity, at the
Experimental Diets and Production of rGH. end of experiment. All data were analyzed by two-
We used 3 types of experimental feed, each with way ANOVA using SPSS statistical software with P
a different proportion of protein; i.e. 21, 28, and > 0.05. Daily growth rate (SGR) was calculated by
34%. Gross energy (GE) for each type of feed the equation:
was the same, approximately 4.200 kcal GE/kg of t Wt
1 is shown in Table 1. rGH
feed. Feed formulation SGR = � �W0 − 1� xX 100%
SGR = 100%
production was performed using Escherichia coli
BL21 containing pCold-1/rElGH vector expression. SGR = daily growth rate; Wt = Average weight
Bacterial culture and rGH collection methods were of an individual at the end of the rearing period
performed as described by Alimuddin et al. (2010). (g); Wo = average weight of an individual at the
beginning of the rearing period (g); t = length of
Table 1. Formulation and chemical composition of the experimental
diets time maintenance (days).
Feed conversion ratio (FCR) was calculated
Protein diets (%) using the equation: FCR = [P/((Wt + Wm)-Wo)]
Ingredients (%)
21 28 34 X100; [FCR = feed conversion ratio; P = amount of
Fish meal 10.00 19.00 26.00 feed given during rearing (g); Wt = biomass of fish
Soybean meal 10.00 23.00 39.00 at the end of the rearing period (g); Wo = biomass of
Pollard 73.00 51.00 28.00
Fish Oil 2.00 2.00 2.00
fish at the beginning of rearing (g); Wm = Weight of
Tapioca 3.00 3.00 3.00 fish that died during rearing (g)]. Protein and lipid
Vitamin and mineral mix 2.00 2.00 2.00 retention were calculated based on Takeuchi (1988).
Proximate (% dry matter) and gross energy (GE) The liver was removed from each fish and weighed
Crude protein 21.28 28.19 34.29 for calculation of the hepatosomatic index (HSI =
Lipid 6.81 6.03 6.14 100 x liver weight/body weight).
Mineral matter 6.94 8.59 10.66 Measurement of Parameters. Analysis of total
Crude fibre 3.85 6.18 5.33
Carbohydrate* 61.12 51.02 43.58
ammonia excretion was determined by measuring
GE (kcal/kg)** 4,337.76 4,236.78 4,284.19 total ammonium nitrogen (TAN) in the culture media
*Carbohydrate = Dry matter – (Crude protein + Lipid + Crude
using a test kit NH3/NH4+ (API © Mars Fishcare North
fibre + Mineral matter); **GE = Gross energy protein 5.6 America Inc., USA). Initial measurement was carried
kcal/g, fat 9.4 kcal/g, carbohydrate 4.1 kcal/g (Watanabe 1988). out after the fish had fasted for 24 h and the aquaria
14 BUDI ET AL. HAYATI J Biosci

were filled with new water (0.19 + 0.08 mg TAN/L).

Table 2. Biomass gain, specific growth rate (SGR), feed consumption, feed convertion ratio (FCR), wet protein body content, protein retention, wet lipid body content, and lipid retention of the giant

Lipid retention (%)

This was performed 24 h after the end of the fish

123.40 + 8.99ab
121.63 + 1.73ab
92.06 + 4.05c

130.46 + 5.50a
88.31 + 3.51c
116.38 + 3.83b
were fed to satiation. TAN was calculated for each

P<0.000
P<0.000
P<0.035
0.066
unit of feed consumed per fish biomass, as modified

*Non rGH is the dietary without rGH supplementation; **Different superscript letters in the same column indicate significant different at P < 0.05 based on Duncan’s multiple range test.
from Suprayudi et al. (2014). Blood glucose levels
were measured by enzymatic colorimetric method
using a liquicolor GLUCOSE test (Human mbH,
Germany). A complete proximate analysis of each

Wet lipid body

8.47 + 0.25bc
8.65 + 0.53b

7.41 + 0.09d
10.05 + 0.25a
8.04 + 0.09c

6.64 + 0.22e
content (%)
fish was carried out on the first and last day of the

P < 0.000
P < 0.000
P < 0.035
0.066
experiment according to the methods of Takeuchi
(1988). Blood triglycerides were measured via
enzymatic colorimetric test with lipid clearing
factor (LCF) using liquicolormonoTRIGLYCERIDES
(Human mbH, Germany). Liver and musle glycogen

Protein retention (%)

gourami (Osphronemus goramy) after feeding with different protein levels and supplemented with recombinant fish growth hormone (rGH)
was measured using the procedure in Wedemeyer and

49.40 + 5.07ab
44.56 + 0.78b
55.39 + 4.79a
54.67 + 2.22a
53.47 + 1.25a
37.38 + 3.05c

P < 0.000
P < 0.001
P < 0.008
Yasutake (1977). Enzyme activity, such as protease,

0.776
amilase, and lipase, were measured according to
Bergmeeyer and Grassi (1983), Worthington (1993),
and Borlongan (1990).

RESULTS
Wet protein body

17.45 + 0.10bc

18.08 + 0.57ab
18.14 + 0.34a
18.50 + 0.42a

16.87 + 0.34c
18.67 + 0.23a
content (%)

P < 0.043
P < 0.001
P < 0.001
Our study results showed that rGH supplemen-

0.171
Parameters

tation of fish diets resulted in higher growth than

diets without rGH supplementation, at each protein
level tested (Table 2). The highest biomass gain,
SGR, and feed consumption, and the lowest FCR
were found in the group given a 34% protein diet
1.66 + 0.09b

1.20 + 0.05d
1.89 + 0.09a

1.35 + 0.04c
1.34 + 0.10c
1.35 + 0.02c

P < 0.000
P < 0.001
P < 0.064
supplemented with rGH. The lowest biomass gain,

0.017
FCR

SGR, and feed consumption, with the highest

FCR were found in the group given a 21% protein
diet without rGH supplementation. Furthermore,
Feed consumption (g)

biomass gain, SGR, feed consumption, and FCR in

356.86 + 39.10abc

405.95 + 27.419a
338.70 + 13.48bc
399.59 + 41.53ab
392.14 + 17.00ab
301.54 + 40.44c

the group given a 28% protein diet supplemented

P < 0.009
P < 0.014
P < 0.407
7.544

with rGH was comparable to that of the control

group receiving a 34% protein diet without rGH
supplementation. In addition, biomass, SGR,
feed consumption and FCR responses showed no
interaction between dietary protein levels and rGH
1.93 + 0.05d

2.37 + 0.04b
2.32 + 0.06b
1.56 + 0.13e

2.13 + 0.08c

2.59 + 0.03a

P < 0.000
P < 0.000
P < 0.262
SGR (%)

supplementation; but each of them independently to

0.017

affect are parameters (Table 2).

We found a significant correlation between dietary
protein levels and rGH supplementation, and protein
159.17 + 13.92e**

and lipid retention, and also protein and lipid body

Biomass gain (g)

214.88 + 12.31d

298.88 + 10.54b
290.33 + 14.93b
339.37 + 9.69a
250.36 + 11.59c

content. There was an interaction between dietary

P < 0.000
P < 0.000
P < 0.854
2.898

protein levels and rGH supplementation. Protein and

lipid body content and retention in rGH treatment
groups at all dietary protein levels were higher than
that of the groups at corresponding dietary protein
Non rGH*

levels who did not receive rGH supplementation,

Non rGH

Non rGH

Two-way ANOVA
Dietary protein

PxR (Interaction)
Standard Error
Protei diets (P)

except for the groups receiving a 28% protein diet

rGH

rGH

rGH
levels

(Table 2).
rGH (R)

We found that HSI was inversely correlated to

21%

28%

34%

dietary protein levels, but that rGH supplementation

Vol. 22, 2015 Growth of Giant Gourami Feeding with Recombinant Growth Hormone �������
15

significantly increased the HSI value of groups at

Table 3. Hepatosomatic index (HSI), total ammonia nitrogen (TAN) excretion, liver glycogen, muscle glycogen, amilase activity, lipase activity, and protease activity of the giant gourami (Osphronemus
each dietary protein levels (Table 3). Furthermore,

(U/mg substrate)
0.019 + 0.004b

0.018 + 0.006b
0.023 + 0.003b
0.024 + 0.008b
0.019 + 0.002b
0.040 + 0.004a
the HSI of the group receiving the rGH supplemented

*Non rGH refers to the diet without rGH supplementation; **Different superscript letters in the same column indicate significant differences at P < 0.05 based on Duncan’s multiple range test.
P < 0.013
P < 0.009
P < 0.002
Protease

0.001
28% protein diet was comparable with the HSI of the
group receiving the 34% protein diet without rGH
supplementation. In this study we did not find that
dietary protein levels and rGH supplementation had
any influence on each other, nor on the effect of either
factor in regards HSI. We found a significant increase

(U/mg substrate)
7.71 + 0.47bc

6.71 + 0.16bc

7.66 + 1.32bc
8.47 + 1.57b
12.01 + 0.91a

6.16 + 0.89c
in TAN excretion with the increase of dietary protein

P < 0.000
P < 0.007
P < 0.004
Lipase

0.237
levels (Table 3). The rGH suplementation decreased
TAN excretion, except in the group receiving an
rGH supplemented 28% test diet compared to the
control group without rGH supplementation. As
with HSI, we found no evidence that dietary protein
levels and rGH supplementation worked in any

(U/mg substrate)
2.80 + 0.08b

2.91 + 0.27b
2.77 + 0.29b
4.22 + 0.58a
2.10 + 0.02c
2.22 + 0.06c
way to alter the effect that each factor would have

P < 0.000
P < 0.005
P < 0.001
Amilase

0.0.068
independently on TAN excretion. TAN excretion in
the group given the rGH-supplemented 28% protein
diet was comparable to the TAN excretion measured
in the group given the 34% protein diet without the goramy) after feeding with different protein levels and supplemented with recombinant fish growth hormone (rGH)
rGH supplementation, while TAN excretion in the
group that received the rGH-supplemented 21% Muscle glycogen
(mg/g sample)

0.53 + 0.11bc
0.91 + 0.37b

0.82 + 0.10b
2.26 + 0.53a
0.23 + 0.01c

0.27 + 0.06c
Parameters

protein diet was comparable to that measured in the

P < 0.000
P < 0.000
P < 0.014
0.065
group that received the 28% protein diet without
rGH supplementation. The lowest TAN excretion
was observed in the group receiving the rGH-
supplemented 21% protein diet supplemented with
rGH, and the highest TAN excretion was found in
the group that received the 34% protein diet without
Liver glycogen
(mg/g sample)

1.98 + 0.47bc
2.83 + 0.66bc
3.12 + 0.37b
7.04 + 0.93a

1.74 + 0.56c
1.86 + 0.66c

rGH supplementation. P < 0.000

P < 0.000
P < 0.001
0.149
The rGH supplementation significantly increased
the levels of blood glucose in treatment groups at
all protein levels, but we did not find that dietary
protein levels and rGH supplementation had any
influence on each other, nor on the effect of either
(mg/g feed/g fish)

factor in regards to blood glucose levels (Figure

0.098 + 0.015bc
0.057 + 0.015d
0.095 + 0.020c

0.172 + 0.016a
0.114 + 0.015b

0.134 + 0.111b
TAN excretion

1). Blood glucose levels in groups receiving rGH-

P < 0.000
P < 0.001
P < 0.387
0.004

supplemented feed (60.67-63.17 mg/dL) were on

average, across all protien levels, 20.67% higher
than those for groups without rGH supplementation
treatments (50.00-54.00 mg/dL). However, triglyceride
levels in all groups, both treated and control, were
1.70 + 0.11b**

comparable, with the exception of the group receiving

P < 0.000
P < 0.000
P < 0.866
1.28 + 0.02d

1.26 + 0.04d
1.91 + 0.05a

1.48 + 0.13c
1.10 + 0.03e

a 21% protein diet without rGH supplementation,

0.018
HSI

whose triglycerides levels were significantly lower

than all treatments (Figure 2).
Both dietary protein levels and rGH supplemen-
tation significantly affected liver and muscle glyco-
Non rGH*

gen levels (Table 3). Unlike the other measurements

Non rGH

Non rGH

Two-way ANOVA
Dietary protein

PxR (Interaction)
Protein diets (P)

Standard Error

in this study, we did find that dietary protein level and

levels

rGH

rGH

rGH

rGH supplementation had an interactive effect on

rGH (R)

each other, increasing the impact each factor would

21%

28%

34%

ordinarily have independently. Liver and muscle

 16 BUDI ET AL. HAYATI J Biosci

70 a a
a
DISCUSSION
60 b
(mg/dL)

b b
Administration of rGH Increased Fish Appetite.
(mg/dL)

50
Appetite is thought to be influenced by increased
level

40 growth hormone stimulation induced by the hormone

level

ghrelin (Volkoff et al. 2005; Debnanth 2010). The

Glucose

30 effect of rGH supplementation in increasing appetite

Glucose

was apparent in the increased feed consumption of

20
fish reared in this research (Table 2). In others, feed
10 consumption also was affected by feed palatability
related to dietary protein levels (Table 2). Palatability
0 of feed relates directly to its attactiveness to fish,
21% 28% 34% determining whether fish will seek, capture, and
Dietary protein levels
Feed protein level ingest (acceptability) of feed. Palatability of feed
Figure 1. Glucose level in blood (mg/dL) of giant gourami relating to nutrient content, especially free amino
(Osphronemus goramy) juvenile after feeding with acids such as glycine, alanine, and betaine; as well
different protein levels and enriched fish recombinant as energy of feed (Shamushaki et al. 2007). In this
without
growth rGH suplementation
hormone with rGH
(rGH). The different letters suplementation
above
the bar indicate significant difference (P < 0.05)
research, the increase in palatability of feed occurs
based on Duncan’s multiple Series1 range tests. without
Series1 in the feed with higher protein content.
rGH suplementation, Series2with rGH suplementation.
Series2
Food intake will affect the activity of digestive
enzymes. Digestive activity is influenced by the
600 substrate directly and by growth hormone (GH)
indirectly (Debhnant 2010; Mataruga et al. 2012;
500 a Irmawati et al. 2012). This was supported by our
a
results: dietary protein levels and rGH administration
Triglycerides level (mg/dL)

a a
400 a triggered digestive enzyme activity, with a trend
that enzyme activity (protease, amylase, and lipase)
34% of lower dietary protein levels treatments were the
28% 300
34%
b same compared to higher dietary protein levels
treatments; rGH giving effect an increase in enzyme
200
activity at 21% protein feed (Table 3).
The increase in blood glucose in groups receiving
100 rGH supplementation was comparable at all dietary
protein treatment levels (Figure 1); protein
0 composition of feed did not seem to affect blood
21% 28% 34% glucose levels. Glucose functions as a source of
Dietary protein levels energy (Hemre et al. 2002). Blood glucose levels
Figure 2. Triglycerides level in blood (mg/dL) of giant gourami were higher in the groups receiving rGH treatments
(Osphronemus goramy) juvenile after feeding with than in those without, regardless of protein levels,
different protein levels and enriched fish recombinant
growth hormone (rGH). The different letters above
and we measured higher energy availability in
the bar indicate significant difference (P < 0.05) the blood after the fish in these groups after they
Series1
based on Duncan’s multiple range tests. Series1 without fasted for 24 h. The high blood glucose levels were
rGH suplementation, Series2with rGH suplementation.
Series2 followed by an increase in liver and muscle glycogen
without rGH suplementation
with rGH suplementation (Table 3). This is consistent with the findings of
glycogen levels of fish fed rGH supplemented 21% Kersten (2001), that unused glucose will be stored
protein diets were higher than those in fish fed as glycogen and the excess will be converted into
21% protein diets without rGH supplementation triglycerides. We confirmed this process in our
treatments. The rGH supplementation in 21% protein finding of an increase in triglyceride levels in groups
diets treatment significantly increased the activity given rGH supplementation, especially in the group
of amilase, lipase, and protease; however rGH receiving the 21% protein diet (Figure 2). Increased
34%
28% supplementation
34% did not significantly affect this triglycerides were measured in the rGH treated fish,
activity for the 28 and 34% protein treatments. after 24 h fasting, due to endogenous synthesis of
There was an interaction between dietary protein triglycerides derived from glucose (lipogenesis) as
levels and rGH supplementation (Table 3).
Vol. 22, 2015 Growth of Giant Gourami Feeding with Recombinant Growth Hormone �������
17

the result of liver glycogen mobilization and the followed by improved FCR measurements (Table
transport of free fatty acids from adipose tissue to 2 and 3). Increased dietary protein along with rGH
the liver (Groff & Gropper 2000). supplementation, resulted in a decrease in FCR,
Glycogen levels rise initially because of an suggesting more efficient feed utilization. This
incrase in liver volumes (Table 3). Yang et al. (2002) indicates that rGH supplementation can improve
similarly saw increases in hepatosomatic index the efficiency of protein and fat utilization to
(HSI) due to the accumulation of glycogen in the increase fish growth. The role of growth hormone
liver of silverperch (Bidyanus bidyanus). The rise in improving feed efficiency has been reported in
in HSI is also related to increased body fat content salmon (Devlin et al. 2004), muchloach (Nam et al.
(Table 2 & 3), with HSI and body fat content highest 2004), and tilapia (Kobayashi et al. 2007).
in the treatment group receiving 21% protein feed. Furthermore, all the parameters that have been
This is in line with the Cheng et al. (2006) study of described are direct effects of the growth of juvenile
grouper (Epinephelus coioides) in which increases giant gourami. The results of this study showed
in body fat content, followed by an increase in liver that the growth measures of juvenile giant gourami
fat content, resulted in an increase in the HSI. increased in correspondence to increases in dietary
Increased protein and lipid retention due to protein and to rGH supplementation. Yang et al.
rGH supplementation has been reported in other (2002) suggests that low dietary protein content
studies (Promdonkoy et al. 2004; Haghighi et results in low growth in many species of fish.
al. 2011). The presence of non-protein energy Meanwhile, the success of oral administration of
(carbohydrates and lipids) can reduce the use rGH in boosting fish growth has also been reported
of protein for energy (protein sparring effect) in several previous studies (Promdonkoy et al. 2004;
resulting in increased protein retention and decreased Haghighi et al. 2011). In our study, we found the
ammonia excretion (Suprayudi et al. 2014). In our highest biomass gain and best SGR in the treatment
study, rGH supplementation allowed for increased group receiving the 34% protein diet supplemented
protein retention (Table 2) and decreased TAN with rGH. Biomass gain and SGR in the treatment
excretion (Table 3). This indicates that higher group receiving the 28% protein diet supplemented
protein retention decreases metabolite excretion with rGH was comparable to that measured in the
of nitrogen (in the TAN form) due to the protein group receiving the 34% protein diets without rGH
sparring effect during the production of energy in supplementation.
fish. This is consistent with Perez-Sanchez (2000), Overall, increased dietary protein, as well as
who found higher nitrogen (protein) retention in rGH supplementation each produced significant
fish receiving rGH treatment, indicating that rGH improvements in the growth of juvenile giant
functions to increase the utilization of non-protein gourami, but we found no interaction between
nutrients as a source of energy (protein sparring the effects of these two treatment variables. The
effect). The role of rGH in reducing TAN excretion interaction between dietary protein levels and rGH
has also been reported in studies of transgenic tilapia supplementation instead was evident in parameters
(Kobayashi et al. 2007). On the other hand, an that supporting growth, such as protein and lipid
increase in dietary protein causes TAN excretion to retention, blood triglyceride levels, liver and muscle
increase (Table 3). Previous research on Australian glycogen, and the activity of digestive enzymes.
short-finned eel juveniles (Guo et al. 2012) and In conclusion, the growth and feed utilization
blue fin trevally (Suprayudi et al. 2014) showed of juvenile giant gourami can be increased by
a positive relationship between protein intake and recombinant growth hormone supplementation in
ammonia excretion. This suggests that amino acids different protein levels of the diets.
from protein, deaminated and excreted as ammonia,
are used to produce energy rather than stored for ACKNOWLEDGEMENT
growth, when fish are given protein supplemented
diets (Mohanta et al. 2008). These results also This report is part of thesis of the first author, which
indicate that low levels of dietary protein and rGH funded by BPPDN scholarship award from DIKTI.
supplementation can reduce ammonia pollution Thanks to Denny Wahyudi, Rangga Garnama, and
from aquaculture. Fajar Maulana for assisting in preparation and
The increases in protein and lipid retention supporting in technical implementation of this
we found due to rGH supplementation, were research.
18 BUDI ET AL. HAYATI J Biosci

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