Indonesian Aquaculture Journal, 14 (1), 2019, 31-38

Available online at: http://ejournal-balitbang.kkp.go.id/index.php/iaj

INFLUENCE OF SQUID LIVER MEAL IN MATURATION DIET ON
GONADOSOMATIC INDEX AND GONADAL AMINO ACID CONTENT
OF GOLDEN SPOTTED RABBITFISH, Siganus guttatus
Asda Laining#, Ike Trismawanti, Usman, and Muhammad Hafid Masruri
Research Institute for Coastal Aquaculture
Jl. Makmur Dg. Sitakka No. 129, Maros 90512, South Sulawesi, Indonesia
(Received 16 March 2018; Final revised 9 January 2019; Accepted 9 January 2019)

ABSTRACT
A four months feeding trial was conducted to evaluate the effects of two maturation diets containing either
fishmeal (FM) or squid liver meal (SLM) as the major protein source on growth, gonadosomatic index and
gonadal bio-chemical profiles of golden spotted rabbitfish, Siganus guttatus. The two tested diets were
formulated to be iso-nitrogenous (40%) and iso-lipidic (14%) and supplemented with several micronutrients
specifically for gonadal development. The initial weight of golden spotted rabbitfish used were 352.6 ±
45.0 g and stocked into four of 2 m x 2 m x 2.5 m net cages with 25 fish per cage. The test diets were fed
to the stocks twice a day. After four months feeding, the weight gains (WG) of broodstock fed the two diets
were nearly similar in which FM had WG of 40.1 ± 2.2% and 42.8 ± 2.0% for SLM. The GSI of male fed SLM
diet was similar to that of fed with FM. However, the GSI of female fed FM was 5.6 ± 0.1% significantly lower
(P<0.05) than SLM which was 11.4 ± 0.5%. Furthermore, total amino acid (TAA) in spermatocyte was
significantly (P<0.05) influenced by dietary SLM indicated by its higher TAA (62.4%) compared to TAA
content of FM group (46.1%). Similarly, the TAA content in oocyte was significantly (P<0.05) improved
when fed SLM. The dietary SLM did not affect the TAA content in the fillet of rabbitfish demonstrated by the
similarity of the fillet TAA content in the two groups which was 55.6% for FM and 54.1% for SLM. Dietary
SLM enhanced GSI of female broodstocks and profile of amino acid in the gonad of golden spotted
rabbitfish, Siganus guttatus.
KEYWORDS:

squid liver meal; maturation diet; gonadal amino acid content; gonadosomatic
index; rabbitfish

INTRODUCTION
Golden spotted rabbitfish, Siganus guttatus is considered to be a promising candidate for marine aquaculture development by virtue of its herbivorous/omnivorous feeding habits and consequent ability to
feed low on the aquatic food chain (Duray, 1998; Tacon
et al., 1990). In nature, rabbitfish feed on algae, seaweed, and other aquatic plants (Parazo, 1990; Bariche,
2006; Jaikumar, 2012) and in captivity can readily to
accept artificial feeds, making them feasible to be
grown on commercial scale (Tacon et al., 1990; Laining
et al., 2017a).

#

Correspondence: Research Institute for Coastal Aquaculture
and Fisheries Extension. Jl. Makmur Dg. Sitakka No. 129,
Maros 90512, South Sulawesi, Indonesia
Phone: + 62 411 371544
E-mail: asdalaining@yahoo.com

Although rabbitfish has several attributes to support their suitability for aquaculture (Saoud et al.,
2008), seed production of the species has not been
successful so far (Avila & Juario, 1987; Rachmansyah
et al., 2007). In addition, commercial diet for maturation stage of rabbitfish so far has not been available
in Indonesia (Laining et al., 2015). Therefore, it is
necessary to develop a broodstock diet to produce
good quality of the seed.
Reproductive performances of fish depend principally on broodstock nutrition (Callan et al., 2012).
Function of various components for broodstock diets such as protein (Tandler et al., 1995), essential
fatty acid (Coman et al., 2011), vitamin E dan C (Binh
et al., 2012; Koshio, 2010), and carotenoid (VasalloAgius et al., 2002; Laining et al., 2017b) have been
intensively studied for several species. The role of
protein and the requirement of specific amino acids

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31

Influence of squid liver meal in maturation dieton gonadosomatic ..... (Asda Laining)

for the production of good-quality eggs have been
better established in marine species (Moran et al.,
2007). Moreover, diet with balanced essential amino
acids improved vitellogenin synthesis in gilthead
seabream (Tandler et al., 1995) while a low protein
and high calorie diet caused a reduction reproductive performance in red sea bream (Watanabe & Kiron,
1995) and seabass (Cerda et al., 1994).
Utilization of certain feed ingredients has been
reported to improve the quality of fish eggs. For instances, squid meal increases the eggs quality of red
seabream, Pagrus major (Watanabe et al., 1991),
gilthead sea bream, S. aurata (Fernandez-Palacios et
al., 1997) and striped jack (Vassallo-Agius et al., 2001)
probably due to its good protein quality and high
content of phospholipid and cholesterol (Watanabe
et al., 1991; Hertrampf & Piedad-Pascual, 2000). In the
market, the are two types of squid meals which are
premium squid meal and squid liver meal (SLM). Premium squid meal is made only from the muscle of
the squid while SLM is produced from whole body of
the squid including gut content (viscera). In order to
determine the influence of squid liver meal (SLM) on
growth, gonadosomatic index (GSI) and bio-chemical profiles of oocyte and spermatocyte, two experimental diets were fed to matured golden spotted
rabbitfish (Siganus guttatus).
MATERIALS AND METHODS
Squid Liver Meal And Experimental
Maturation Diets
Several protein sources were utilized to formulate the two maturation diets with fish meal and squid
liver meal as the tested ingredients (Table 1). All these
protein sources were locally purchased excluding the
Korean SLM. Among the protein sources, fish meal
contained the highest protein content of approximately 65%. The SLM used in this experiment contains protein content approximately 43%. As the car-

bohydrate sources, wheat four was included in the
two maturation diets at the same level of 8%, while
sorghum was only added in one diet to adjust its
protein level.
The experiment consisted of two maturation diets containing either fish meal (FM) or squid liver
meal (SLM) as the major protein source at the level
of 45%, respectively. Each of the diet was still formulated to contain either fish meal or SLM at the minimum level of 5%. The two tested diets were formulated to be iso-nitrogenous (40%) and iso-lipidic (14%)
and supplemented with several micro-nutrients specifically for gonadal development including vitamin
C, vitamin E, and carotenoid mixture (Table 2). In order to meet the required protein level of SLM diet, it
was also incorporated with mysid meal at level of
20%. All dry ingredients were mixed thoroughly and
oil containing lipid solubled-vitamins and carotenoid
was added and then mixed again for 30 minutes. Further, water was added at level of 350-400 g/kg of the
dry ingredients (Laining et al., 2017a). The mixture
was then passed through a pelletizer (Hiraga, Co. Ltd,
Kobe, Japan) and dried in oven. All test diets were
stored in air-tight polyethylene bags in a freezer (20°C) until used.
Initial weight of wild golden spotted rabbitfish
used were 352.6 ± 45.0 g and stocked into four of 2
m x 2 m x 2.5 m net cages with 25 fish per cage. The
test diets were fed to the broodstocks twice a day in
the morning at 08:00 and in the afternoon at 16:00.
The feeding trial lasted for four months.
Sample Collection, Bio-Chemical Analysis,
and Statistic
At the end of the feeding trial, three maturing
fishes for both male and female were collected from
each cages and sacrificed for measuring the GSI and
HSI. After weighing, the gonad samples (oocytes and
spermatocytes) were freeze-dried and kept at -20°C

Table 1. Proximate composition (% dry basis) of feed ingredients used as protein
and carbohydrate sources for the experimental maturation diets
Ingredients
Fish meal
Mysid meal
Soybean meal
Squid liver meal
Wheat flour
Sorghum

Crude protein

Lipid

Ash

Fibre

NFE

65.0
53.0
40.5
43.0
12.0
8.4

7.4
4.0
17.5
10.9
0.3
5.0

20.8
NA*
5.8
10.5
6.2
2.7

1.2
2.7
8.6
12.3
4.6
2.7

5.6
27.6
23.3
76.9
81.2

Note: * NA: not analyzed

32

Copyright @ 2019, Indonesian Aquaculture Journal, p-ISSN 0215-0883; e-ISSN 2502-6577

Indonesian Aquaculture Journal, 14 (1), 2019, 31-38

Table 2. Formulation and proximates composition of experimental
maturation diets
Ingredients (%)

Experimental maturation
FM

SLM

Fish meal
Mysid meal
Soybean meal
Squid liver meal
Wheat flour
Sorghum
Fish oil
Olive oil
Vitamin mix
Mineral mix
Organic mineral
Carotenoid mixture*
Lecithin
Aquasterol
Stay-C
Vitamin E
CMC

45
5
5
5
8
13
5
3
3
3
1
0.77
0.2
0.5
0.1
0.04
2.39

5
20
5
45
8
0
3
3
3
3
1
0.77
0.2
0.5
0.1
0.04
2.39

Proximate composition (%, dry basis)
Lipid
Crude protein
Fiber
Ash

14.0
40.2
3.7
**NA

14.4
40.9
3.7
NA

Note:

* Astaxanthin source 0.2%, cantaxanthine source 0.07%, and Spirulina powder
0.5%; ** NA: not analyzed

until analysis. Dissected fish were then filleted to take
the muscle, oven-dried and kept at -20°C until chemical analysis. Bio-chemical analysis for samples (feed,
oocyte, spermatocyte, and muscle) were performed
for proximate analysis according to AOAC International (2005). Briefly, moisture was analyzed after drying the samples at 105°C for 16 hours using an oven
(Memmert, Germany). Ash was analyzed using a muffle
furnace at 550°C (Barnstead, Thermolyne, CA, USA).
Crude protein was determined according to microKjeldahl procedure (digestion, distillation, and titration). Lipid was extracted using chloroform and
methanol. Furthermore, analysis of total amino acid
was performed with HPLC (Shimadzu 20A, Tokyo, Japan).

liver/body weight). All data obtained were statiscally
analyzed using Independent t-test with software SPSS
(version 21; SPSS, Inc., Chicago, Illinois, USA). Differences were considered significant at P<0.05.

Variables obser ved were weight gain (WG),
gonadosomatic index (GSI), hepatosomatic index
(HSI), and amino acid profiles of gonads and fillet of
both male and female broodstocks. GSI (%) was calculated as GSI= 100 x (weight of gonad/body weight)
and HSI (%) was calculated as HSI= 100 x (weight of

Growth response observed in the present study
implied that SLM has no benefit on gowth when fish
are given at maturation stage. Unlikely when fish fed
during the growth phase, a positive correlation was
found between the growth performance of juvenile
Japanese seabass (Lateolabrax japonicus) and the supple-

RESULTS AND DISCUSSION
After four months feeding, the weight gains (WG)
of broodstock fed the two diets were nearly similar
in which FM had WG of 40.1 ± 2.2% and 42.8 ± 2.0%
for SLM. The growth of fish from the two groups
increased slowly during the four months culture as
illustrated by Figure 1. The slow growth of the fish
in both groups indicated that the fish were in the
maturation stage or the fish has been categorized as
a broodstock.

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33

Influence of squid liver meal in maturation dieton gonadosomatic ..... (Asda Laining)

Individual weight (g)

600
500
400
300
200
FM
SLM

100
0
1

2

3

4

5

Days of culture

Figure 1.

Pattern of weight increment of golden spotted rabbitfish
S. guttatus fed two maturation diets.

mentation of squid viscera meal (SVM), indicating that
SVM can stimulate juvenile to grow (Mai et al., 2006).
Liang et al. (2000) also reported that SVM supplementation increased the growth of three marine species
Chrysophrys major, Fugu rubripes and Moroul satatilis.
Kader et al. (2012) revealed that fermented soybean
meal combined with squid by-product blend stimulated the growth, feed utilization, and health of Japanese flounder, Paralichthys olivaceus. The very slow
growth of the rabbitfish fed the two diets found in
the present study might be the result of a genetic
predisposition (maximum size achieved for older fish)
(Mejri et al., 2017). For instances, tiger shrimp at age
of 11 months grew faster (170% of WG) than those
with age of 16 months (30% of WG) (Laining et al.,
2017b).
Table 3 presents the GSI of male fed SLM diet was
similar to that of fed with FM which was 7.5 ± 0.2%
and 8.1 ± 0.3%, respectively. The GSI of female fed
FM diet was 5.6 ± 0.1% significantly lower than GSI
of female fed SLM which was 11.4 ± 0.5%. The significant effect of SLM on GSI of female may imply the
effect of SLM on fecundity which was not measured
in the present study. Recent studies evaluating the
SLM or other types of squid meal on reproductive
performances are not readily available, and therefore
difficult to compare with findings of the present
study. However, Fernandez-Palacios et al. (1997) revealed that fat-insoluble fraction of squid meal improved the egg quality of gilthead sea bream (Sparus
aurata) broodstock in regard to fecundity produced
per kilogram of female and percentages of viable and
fertilized eggs. Even though, the SLM significantly
enhanced the GSI of female spawners, no significant
effect (P>0.05) was observed on GSI of male
broodstocks.
34

Proximate composition of oocyte and spermatocyte are presented in Table 4. No differences were
found on the macro-nutrient content of oocyte for
both goups of fish fed FM and SLM diets. In term of
spermatocyte, the crude protein, and lipid content
were similar for both goups of fish fed SLM and FM.
However, ash content was slightly increased in fish
fed SLM. Furthermore, crude protein content of spermatocyte was higher than in oocyte while the lipid
content was lower in spermatocyte than in oocyte.
These may indicated that lipid is more necessary for
oocyte than in spermatocyte. Nutrient in eggs depend primarily on broodstock nutrition (Callan et al.,
2012), while the approppriate biochemical composition in eggs is necessary to meet the development
requirements of embryonic and larval stages (Araujo
et al., 2012). The high protein content in both oocyte and spermatocyte suggested that protein is highly
required for gonadal development of rabbitfish. Protein is the most abundant component of the nutrients contained in the egg of many of the fish species
studied (Watanabe & Vasallo-Agius, 2003).
Furthermore, total amino acid (TAA) in spermatocyte was positively influenced by dietary SLM indicated by its higher TAA (62.4%) compared to TAA
content of FM group (46.1%) as presented by Table 5.
The same pattern was also observed in oocyte where
the TAA content in oocyte was improved when fish
fed SLM rather than FM diet (49.3% vs 30.1%, respectively). The significant higher TAA both in oocyte and
spermatocyte of fish fed SLM than those fed with FM
diet indicated that protein of SLM may be more available and more benefit for gonadal development of
rabbitfish compared to fish meal. The dominant
amino acid in spermatocyte of rabbitfish found in SLM
whose content was more than 5% of TAA were as-

Copyright @ 2019, Indonesian Aquaculture Journal, p-ISSN 0215-0883; e-ISSN 2502-6577

Indonesian Aquaculture Journal, 14 (1), 2019, 31-38

Table 3. GSI (%) and HSI (%) of female and male stocks of golden spotted
rabbitfish fed two maturation diets
Diets

Female

Male

GSI

HSI

GSI

HSI

FM

5.6 ± 0.1 a

2.0 ± 0.0 a

7.5 ± 0.2 a

1.5 ± 0.0a

SLM

11.4 ± 0.5 b

3.5 ± 0.2 b

8.1 ± 0.3 a

1.7 ± 0.0a

Description: All values are mean ± SD. Different superscript letters within each group
indicate significant different mean (P<0.05)

Table 4. Proximate composition (% dry basis) of oocyte and spermatocyte of golden
spotted rabbitfish fed two maturation diets
Nutrients

Oocyte

Spermatocyte

FM

SLM

FM

SLM

Lipid

8.4 ± 0.1 a

8.6 ± 0.4 a

2.6 ± 0.4 a

3.1 ± 0.3a

Crude protein

68.5 ± 0.6 a

68.5 ± 0.7 a

72.7 ± 2.3 a

76.7 ± 0.6 a

Ash

12.8 ± 1.4 a

10.9 ± 0.1 a

11.6 ± 0.4 a

14.2 ± 0.5 b

Description: All values are mean ± SD. Different superscript letters within each group indicate
significant different mean (P<0.05)

partic acid, glutamic acid, glycine, arginine, and leucine. In oocyte, the dominant amino acid content (>
5%) was only glutamic acid and other amino acids including aspartic acid, arginine, alanine, valine, isoleucine, leucine, and lysine had moderate level between
3%-5% of TAA. According to Mejri et al. (2017) amino
acid leucine, lysine, valine, isoleucine, alanine, aspartic acid, serine, and glutamic acid were important in
eggs and larvae of walleye (Sander vitreus), with the
highest values reported for lysine and leucine in essential amino acid fraction. In gilthead seabream egg,
amino acid composition of vitelline envelope proteins is characterized by a high content of proline
and glutamic acid and relatively low content of cysteine (Hyllner et al., 1995).
The dietary SLM did not significantly (P>0.05)
affect the TAA content in the fillet of golden spotted
rabbitfish when fed at maturation stage demonstrated
by the similarity of the fillet TAA content in the two
groups of fish which was 55.6% for FM and 54.1% for
SLM (Table 6). The individual amino acid in fillet was
also not influenced by dietary SLM. These pattern
was different from amino acid profile in gonads in
which dietary SLM significantly improved most of their
amino acid contents.
Table 7 presents the proximate composition of
liver after four months feeding trial. There was no

clear effect of the dietary treatments on the nutrient
content in the liver. The protein content was approximately 45% in both groups of fish fed FM and SLM
diet, respectively. The liver lipid content was relatively low which was 4.1% for FM and 2.0% for SLM.
Based on GSI values and amino acid content in
both oocytes and spermatocytes, reproductive performance of golden spotted rabbitfish could be improved by using LSM as the major dietary protein
source. Concentration of TAA in spermatocytes of
fish fed LSM diet increased approximately 35% and
the improvement was more pronounced observed in
oocytes which its TAA increased 63.8% compared to
fish fed FM diet. According to Penaflorida (1989),
amino acid profile of squid meal is well balanced with
index is high (0.96) similar to white fish meal (0.96)
and shrimp meal (0.98). Besides its balanced amino
acid profile, rabbitfish may have ability to absorb nutrient in SLM more efficiently than FM. Although determination of nutrient digestibility coefficient of
SLM was not carried-out for this present experiment,
squid meal has high protein, and dry matter digestibility for tiger prawn which is 96.0% and 85.5%,
repectively (Hertrampf & Piedad-Pascual, 2000). This
indicates that tiger shrimp can efficiently utilise dietary LSM. However, the positive effects of dietary
LSM on amino acid content was not observed in fil-

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Influence of squid liver meal in maturation dieton gonadosomatic ..... (Asda Laining)

Table 5. Amino acid profile (%, dry matter) of oocyte and spermatocyte of golden
spotted rabbitfish fed two maturation diets
Spermatocyte

Amino acid

Oocyte

FM

SLM

FM

SLM

Aspartic acid
Glutamic acid
Serine
Histidine
Glycine
Threonine
Arginine
Alanine
Tyrosine
Methionine
Valine
Phenylalanine
Isoleucine
Leucine
Lysine

3.87 ± 0.06 a
7.43 ± 0.03 a
1.84 ± 0.08 a
1.10 ± 0.17 a
3.78 ± 0.08 a
2.20 ± 0.29 a
5.42 ± 0.14 a
3.50 ± 0.03 a
1.61 ± 0.06 a
1.12 ± 0.02 a
2.84 ± 0.07 a
1.89 ± 0.06 a
2.20 ± 0.23 a
4.12 ± 0.09 a
3.16 ± 0.05 a

5.49 ± 0.12 b
9.81 ± 0.04 b
2.29 ± 0.08 b
1.32 ± 0.06 a
5.12 ± 0.03 b
2.88 ± 0.15 a
8.53 ± 0.01 b
4.02 ± 0.06 b
2.18 ± 0.02 b
1.45 ± 0.06 b
3.71 ± 0.14 b
2.48 ± 0.08 b
2.89 ± 0.11 a
5.61 ± 0.14 b
4.60 ± 0.14 b

3.30 ± 0.14 a
5.01 ± 0.01 a
1.12 ± 0.15 a
0.84 ± 0.04 a
3.02 ± 0.05 a
1.53 ± 0.02 a
2.49 ± 0.05 a
1.73 ± 0.02 a
0.28 ± 0.03 a
0.88 ± 0.03 a
1.88 ± 0.03 a
1.55 ± 0.03 a
1.70 ± 0.06 a
2.46 ± 0.01 a
2.31 ± 0.03 a

4.55 ± 0.08 b
6.96 ± 0.07 b
2.91 ± 0.03 b
1.32 ± 0.01 b
2.14 ± 0.02 b
2.59 ± 0.01 b
3.84 ± 0.02 b
3.53 ± 0.08 b
2.16 ± 0.03 b
1.10 ± 0.02 b
3.60 ± 0.02 b
2.68 ± 0.02 b
3.20 ± 0.03 b
4.74 ± 0.02 b
3.99 ± 0.16 b

Total

46.09 ± 1.19 a

62.4 ± 0.11 b

30.1 ± 0.18 a

49.31 ± 0.08 b

Description:

All values are mean ± SD. Different superscript letters across columns within each
group indicate significant different mean (P<0.05)

Table 6. Amino acid profile (%, dry matter ± SD) of female stock fillet
of golden spotted rabbitfish fed two maturation diets
Parameter

Fillet FM

Fillet SLM

Aspartic acid

5.58 ± 0.08a

5.91 ± 0.14 a

Glutamic acid

8.64 ± 0.03a

9.38 ± 0.23 a

Serine

2.25 ± 0.09a

2.40 ± 0.16 a

Histidine

5.91 ± 0.06a

1.29 ± 0.04 a

Glycine

2.99 ± 0.18a

3.16 ± 0.13 a

Threonine

2.73 ± 0.03a

2.87 ± 0.18 a

Arginine

3.61 ± 0.04a

3.84 ± 0.11 a

Alanine

3.44 ± 0.03a

3.66 ± 0.28 a

Tyrosine

1.87 ± 0.16

a

1.94 ± 0.06 a

Methionine

1.57 ± 0.17a

1.74 ± 0.06 a

Valine

2.91 ± 0.27a

3.08 ± 0.20 a

Phenylalanine

2.26 ± 0.09a

2.40 ± 0.71 a

l-leucine

2.71 ± 0.05a

2.85 ± 0.35 a

Leucine

4.33 ± 0.13a

4.64 ± 0.42 a

Lysine

4.74 ± 0.08a

4.96 ± 0.21 a

Amino acid total

55.37 ± 0.34 a

54.17 ± 0.12 a

Description: All values are mean ± SD. The same superscript letters across
columns indicate no significant different mean (P>0.05)

36

Copyright @ 2019, Indonesian Aquaculture Journal, p-ISSN 0215-0883; e-ISSN 2502-6577

Indonesian Aquaculture Journal, 14 (1), 2019, 31-38

Table 7. Proximate composition (% dry basis ± SD) of liver of rabbitfish S.
guttatus fed two maturation diets
Nutrient
(%)
Lipid
Crude protein
Fiber
Ash

let, possibly indicated that certain amino acid level in
gonads were more sensitive to be affected by dietary
SLM than by dietary FM.
CONCLUSION
Dietary SLM enhanced GSI of female broodstocks
of rabbitfish from 5.6% to 11.4% but did not affect
the GSI of male broodstock. Total amino acid content significantly improved by dietary LSM from
30.1±0.18 to 49.31±0.08% in oocyte and from
46.09±1.19 to 62.4±0.11% in spermatocyte of golden
spotted rabbitfish, Siganus guttatus.
ACKNOWLEDGEMENTS
The authors thank to the Australian Center for
International Agricultural Sciences (ACIAR) for funding this research through Project FIS/2007/124 Diversification of smallholder coastal aquaculture in Indonesia
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4.1 ± 0.4
45.1 ± 2.5
NA
4.2 ± 0.1

2.0 ± 0.3
44.9 ± 0.4
NA
3.9 ± 0.4

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