Journal Pre-proof

Feed intake stimulation of juvenile spotted scat (Scatophagus

argus Linnaeus, 1766) using dietary seaweed supplementation
(Ulva sp.)

Monsuang Yangthong, Jirayuth Ruensirikul

PII: S0044-8486(19)32786-3
DOI: https://doi.org/10.1016/j.aquaculture.2020.735626
Reference: AQUA 735626

To appear in: Aquaculture

Received date: 20 October 2019

Revised date: 14 June 2020
Accepted date: 15 June 2020

Please cite this article as: M. Yangthong and J. Ruensirikul, Feed intake stimulation
of juvenile spotted scat (Scatophagus argus Linnaeus, 1766) using dietary seaweed
supplementation (Ulva sp.), Aquaculture (2020), https://doi.org/10.1016/
j.aquaculture.2020.735626

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Feed Intake Stimulation of Juvenile Spotted Scat (Scatophagus argus Linnaeus,

1766) Using Dietary Seaweed Supplementation (Ulva sp.)

Monsuang Yangthonga,* and Jirayuth Ruensirikulb

a
Program of Fishery Science and Aquatic Resources, Faculty of Agricultural

Technology, King Mongkut’s Institute of Technology Ladkrabang, Prince of Chumphon

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Campus, Chumphon, 80160 Thailand

b pr
Coastal Aquaculture Research and Department Regional Center 6, Songkhla, 90000
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Thailand
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*
Corresponding author. Tel.: +66 9-9435-2888; fax: +66 7-7591-445.

E-mail address: ymonsuang@hotmail.co.th (M. Yangthong).

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Sample CRediT author statement

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Monsuang Yangthong: Conceptualization, Methodology, Software

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Monsuang Yangthong.: Data curation, Writing- Original draft preparation.

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Jirayuth Ruensirikul : Visualization, Investigation.

Michael Currie : Writing- Reviewing and Editing,

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Abstract

This study appraised the growth performance of juvenile spotted scat when fed diets

supplemented with Ulva sp. Seven isonitrogeneous and isocaloric diets with different

levels of supplementation with Ulva sp. at 0 (control), 5, 10, 15, 20, 25 and 30% were

fed to 420 fish, weighing 0.49 ± 0.11 g. The diets were fed to triplicate groups of fish

twice a day to apparent satiation for 10 weeks. The results showed that fish fed the diets

with 5 and 10% seaweed demonstrated the highest mean WG and SGR (P < 0.05),

f
whereas there were no significant differences in the FCR and PER of fish fed the

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control diet, and 5 and 10% seaweed (P > 0.05). Fish fed the control diet and that

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containing 5% Ulva sp. showed a lower rate of FI than fish fed diets containing 10–30%

(P < 0.05). Moreover, dietary supplementation with seaweed at levels of 5–25% had no
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significant effect (P > 0.05) FCE although it was higher for those diets than for the fish
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fed the control diet and that supplemented with 30% Ulva sp. (P < 0.05). Therefore,

10% Ulva sp. in diet should be applied for spotted scat rearing.
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Keywords: Ulva sp.; spotted scat; feed supplement

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1. Introduction

The spotted scat is a tropical euryhaline species that is a candidate for commercial

coastal aquaculture in this region (Gupta, 2016) because of several interesting

characteristics such as good adaptation to highly fluctuating environments (Barry and

Fast, 1992). Although up to the present, studies in many countries have succeeded in

breeding this fish (Chang, 1997; Ruensirikul et al., 2008; Cai et al., 2010; Khanh et al.,

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2012; Gandhi et al., 2014;), mass seed production of spotted scat is still limited, and the

survival rate of juvenile scat reared from hatching is variable and appears to be low. In
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addition, slow growth and a long period of weaning have been observed. One of the
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main causes may be an inappropriate nutritional content in food given to fish fry or an
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unattractive artificial diet being fed to them. However up to now there has been a lack

of research into the optimum formulation of feed for spotted scat.

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The seaweed, Ulva, a green alga in the division Chlorophyta, also known by the
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common name, sea lettuce, is common from tropical to polar climates, although strains
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appear to vary from region to region. Ulva has been found to contain high amounts of
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good-quality protein, carbohydrates, vitamins and minerals (Mahae et al., 2014) and is a

good source of many important nutrients and a potent alternative plant nutrient source

for both humans and animals due to its low toxicity and high level of bioactivity

(Peixoto et al., 2016). For instance, a sulfated polysaccharide extracted from the cell-

walls of Ulva has been found to demonstrate antioxidant, antibacterial, and

immunostimulant properties (Shao et al., 2013).

The use of Ulva as a fish-feed ingredient or feed supplement has been reported by a

number of studies both as a means of reducing feed cost and enhancing growth
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performance (Bindu and Sobha, 2004; El-Tawil, 2010; Zinadah et al., 2013; You et al.,

2014; Abdel-Warith et al., 2016; Zhang et al., 2017). However, there have been no

reports of the use of Ulva as a fish feed for spotted scat culture during the early

development of juvenile fish although, seaweed is one of the main items found in the

gut content of this species in its natural habitat (Sivan and Radhakrishnan, 2011). In scat

hatcheries, the period during which young fish adapt to their normal adult diet is usually

long and consequently involves high production costs due to the overuse as feed of the

brine shrimp (Artemia naupli) which is expensive. As a result the feed intake of

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artificial diets is likely to be low.

The feed intake of fish is related to their need for amino acids such as aspartic acid
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and glutamic acid (Carr et al., 1996). Mahae et al. (2014) found that aspartic acid was
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the main non-essential amino acid in cultured-Ulva. However, the biochemical
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composition of Ulva collected from different sites may differ (Ktari, 2017; Selvin et al.,

2011; Yangthong et al., 2009; Yedukondala Rao et al., 2015). Nevertheless, the
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utilization of Ulva as a feed ingredient may benefit fish-feed intake and growth
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performance. Therefore, the research described studied the growth rate and feed intake
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of cultured-spotted scat fry fed a diet supplemented with Ulva sp. to establish the
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optimum level of supplementation for raising this fish.

2. Material and methods

2.1 Diet formulations

Dried Ulva sp. seaweed was obtained from the Trat Coastal Fisheries Research and

Development Centre, Thailand, dried in an electrical oven at 55°C for 48 h and milled

using a herb grinder (Thai Grinder, model NT 500D). The ground materials were kept

in air-tight plastic bottles at 4°C for proximate analysis (AOAC, 1995), while other
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dietary ingredients were purchased from a feedstuff shop and the components were

analyzed and are listed in Table 1. The seven diets formulated contained approximately

39% crude protein. Diet 1 without seaweed served as the control diet, while diets 2, 3, 4,

5, 6 and 7 were formulated such that seaweed replaced proportions of 5, 10, 15, 20, 25

and 30%, respectively of the standard fish meal. Details of the diet formulation and

proximate composition are shown in Table 2. The dietary ingredients were mixed with

water to produce 1.5 mm pellets using a pelleting machine (La Minerva A/E12, Italy).

The dietary pellets were dried in a hot air oven at 70°C and subsequently stored at 4°C.

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[Table 1]

[Table 2]
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2.2 Experimental design
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Juvenile spotted scat (Scatophagus argus) were obtained from the marine fish

hatchery at the Coastal Aquaculture Research and Development Regional Center 6,

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Songkhla, Thailand. The fish were acclimatized to laboratory conditions with self-
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prepared feed for 2 weeks. Four hundred and twenty acclimatized fish, weighing 0.49 ±
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0.11 g were divided into seven groups in three replicates of 20 fish for each group per
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aquarium. Each aquarium (90 × 40 × 45 cm) was filled with 140 L of seawater and

supplied with continuous aeration. The fish were fed to apparent satiation twice a day at

8:30 a.m. and 16:30 p.m. During the experimental period, the salinity and water

temperature were maintained at 5 ppt and 25–30°C, respectively with constant aeration

and a natural photoperiod. The levels of dissolved oxygen, ammonia (NH3 ) and nitrite

(NO 2 ) were maintained and remained at acceptable levels throughout the period of the

experiment. Dissolved oxygen was measured by portable DO meter (WTW Multiline

P4, Germany), ammonia and nitrite were measured using standard test kits (V-unique,
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Better Syndicate Co., Ltd., USA) Every day before feeding, half of the aquarium water

was replaced, and the feces and pellet residues were removed by siphoning. The fish

were weighed at the beginning of the experiment and then bi-weekly for 10 weeks.

Before weighing, the fish were starved for 24 h, allowing the gut to empty.

The growth and biological indices of the fish were established in terms of mean

weight gain (WG), specific growth rate (SGR), feed intake (FI), feed conversion

efficiency (FCE), feed conversion ratio (FCR), protein efficiency ratio (PER), lipid

efficiency ratio (LER), energy efficiency ratio (EER) and survival rate (SR). These

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indices were calculated as follows:

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WG: [final body weight (g) − initial body weight (g)] × 100 / initial body weight (g)
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SGR: [Ln final body weight (g) − Ln initial body weight (g)] × 100 / experimental
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period

FI:
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feed intake g
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initial body weight g final body weight g initial fish uantity final fish uantity e eri ental eriod
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FCE: [(body weight gain (g) / feed intake (g)) × 100]

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FCR: feed intake (g) / weight gain (g)

PER: weight gain (g) / protein intake (g)

LER: weight gain (g) / lipid intake (g)

EER: weight gain (g) / energy intake (kcal)

SR: [final fish quantity / initial fish quantity] × 100

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2.3 Chemical analysis

The seven diets were dried at 80°C before analysis. The proximate composition was

determined according to the standard methods of analysis (AOAC, 1995): dry matter

(DM, 24 h at 100 ± 1°C), ash (3 h at 600°C), crude protein (Nx6.25 Kjeldahl method)

and fat content (petroleum ether 160°C extraction in Soxlet System HT6) crude fibre by

extraction with 0.5 M of H2 SO 4 , and ash by using a muffle furnace at 600°C for 15 h.

The gross energy was measured by the ballistic bomb calorimetric method (AOAC,

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1995). The amino acid content of the diets was determined by HPLC (Waters

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Alliance2695 with heater, Jasco FP2020 fluorescence detector (EX:250 and EM: 395

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nm)) with a Hypersil gold column C18 (4.6×150 mm, 3µm) at 35 C at The Central
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Instrument Facility, Faculty of Science, Mahidol University.
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2.4 Statistical analysis

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All the data were reported as mean  standard deviation (SD) and the variance in the
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data was analyzed using one-way ANOVA with significant differences of mean values

being established using Duncan’s new ulti le range test. Differences were considered

as being significant at P < 0.05.

3. Results

3.1 Amino acid content of diets

The concentrations of amino acid in the experimental diets are shown in Table 3.

The concentrations of essential amino acid in the diets (histidine, isoleucine, threonine,
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proline, lysine and leucine) and non-essential amino acid (glutamic and aspartic acid,

alanine and glycine) were different among the treatments. Supplementation with Ulva

tended to increase the concentrations of amino acids in the diets. The amounts of

glutamic and aspartic acid were higher than any other amino acid and the glutamic and

aspartic acid, alanine, isoleucine, threonine and proline concentrations in the diet

supplemented with Ulva were higher compared to the control diet.

[Table 3]

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3.2 Growth performance

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The growth performance and survival rate during the spotted scat feeding trial are

summarized in Table 4. The study found that the growth performance of fish varied
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with the degree of algal supplementation in the diets. Fish fed 5% Ulva sp.
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demonstrated the highest mean WG and SGR. The WG and SGR of fish fed the control

diet were lower than those of the fish fed with 10–20% supplementation of Ulva sp., but
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for the group fed diets with 25–30% Ulva sp. the WG and SGR were lower than those
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of the fish fed the control diet. However, none of the groups fed different proportions of
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Ulva sp. in their diets showed any significant differences in SR from that of the control
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group.

[Table 4]

3.3 Feed utilization

The feed utilization data for the fish fed the seven diets are set out in Table 5. Algal

supplementation in fish diets was found to stimulate the appetite of the fish as shown in

Figure 1. The mean daily FI showed significant differences (P < 0.05) between the fish

fed each of the seaweed supplemented diets and the control group. Fish fed 10–30% of
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algae demonstrated the highest mean rate of FI (Figure 1) while the intake of even the

fish fed only 5% algae was significantly higher than that of the control group. The FCE

of the fish was also found to vary with the level of algal supplementation in the diets.

Fish fed with 5% algal supplementation demonstrated the highest mean FCE while the

FCEs of the fish fed on the 10–25% algae diets were significantly better than those fed

the control diet and that supplemented with 30% algae. There were no significant

differences in the FCR, PER and LER between the fish fed the control diet and fish

those fed either the 5 and 10% algal supplemented diets, but these had the highest PER

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and LER values, although for LER the group fed a 20%-supplemented diet also showed

a high value, Meanwhile the fish receiving 15–30% algae diets displayed generally
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decreased PER and LER with increasing algal supplementation in their diets. The EER
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was also not significantly different between fish fed the control diet and those fed 5–
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25% algae. However, the EER value was at its maximum for the fish fed the 5% algae

diet then decreased with increasing algal supplementation in the diets.

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[Table 5]
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[Figure 1]
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3.4 Water quality

The water quality parameters measured in the tanks in which the spotted scat fed the

experimental diets were kept for 10 weeks, showed no differences among the

treatments, nor among the SR of the fish. The temperature ranged from 25.00–30.00°C,

the salinity was 0055 parts per thousand, the dissolved oxygen was in a range of – 5050

30.7 mg/L, the nitrite was in a range of 5055 to 003. mg /L, and the total ammonia

content was 0.03–0.94 mg /L.

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4. Discussion

Ulva has the potential to be applied as a raw material for the production of feed for

spotted scat cultivation. In the present study, with Ulva sp. supplementation, the diets

displayed higher protein and fat contents while the moisture and ash contents were

lower than those found in U. rigida collected from the coastal area of the Dardanelle

Strait in Turkey, for which the nutritional values were 0.15, 11.5, 26.4%, protein, ash

and fat, respectively (Kut Güroy et al., 2007). Latuihamallo and Loupatty (2016)

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reported the protein, water, ash and fiber contents of U. fasciata to be 700, 1.7, 25.7 and

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4.8% respectively, Yaich et al. (2011) found protein, fat and ash contents for U. lactuca

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of .0.5, 7.87 and 19.59%, respectively, and Abdel-Warith et al. (2016) found 55005,
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6.08 and 24.29%, respectively. In general, the protein content of Ulva seaweed has been

noted to be between 10 and 26% (Fleurence, 1999).

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Ulva is a macro seaweed which is commonly found in the stomach content of wild
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spotted scat (Gandhi, 2002). This seaweed, should, therefore, be used as a feed

supplement for this variety of fish. It is of interest, however, to consider the appropriate
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level of supplementation that can be applied with this fish and this can be evaluated
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based on fish growth as indicated by WG and the SGR. In the present study,
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supplementing their feed with 5 and 10% Ulva showed benefit in the growth

performance of the fish compared to the non-supplemented feed in the control

treatment. Muangyao et al. (2013) investigated the growth performance of 3.18 ± 5 g

spotted scat fed a diet supplemented with fresh U. intestinalis at levels of 0, 5, 10 and

15%. They found that were no significant differences in weight gain among the groups

fed all the supplemented levels, and the lowest FCR was observed in fish fed a 10%

seaweed-supplemented diet, while the lowest FCR was found in the fish in the (non-

supplemented diet) control group. In the present study, the fish fed the control and 5 or
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10% supplemented diets showed no differences in FCR. Valente et al. (2006) however,

found that this growth parameter was higher with higher levels of Ulva sp.-

supplementation from 0 to 10% in a study of the growth of 4.7 g European sea bass

(Dicentrarchus labrax) for 10 weeks. They found that the weight gain was lower and

the FCR was higher with increased levels of seaweed supplementation while the best

WG and FCR were found in the non-seaweed-supplemented feed group of control fish.

However, the PER of the European sea bass fed all levels of supplementation was not

significantly different, and Ulva could therefore be safely added to the feed for this fish

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at proportions of up to 10%.

In the present study, the PER and LER generally decreased with increases in the
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seaweed content of the fish feed. These parameters were most satisfactory in the fish
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that were fed with 0–10% seaweed levels while the report of El-Tawil (2010), who
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studied 6 levels (5, 5, 10, 15, 20 and 50%) of Ulva sp. feed supplementation in 1.5 g

tilapia (Oreochromis sp.) for 9 weeks, found that fish fed with 5 5 and 15%
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supplementation achieved the best WG and SGR. while the highest FCRs were
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observed in fish fed with 20, 15 and 10% seaweed-supplemented feed.

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Kut Güroy et al. (2007) studied the effect of U. rigida-feed supplementation on

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tilapia (Oreochromis niloticus) at 0, 5, 10 and 15% and found that there were no

differences in the WG and SGR of fish fed with 0, 5 and 10% seaweed-supplemented

feed although these parameters were better than those of fish fed with 15% Ulva-

supplemented feed. However they found that there was no difference in the FI among

all the treatment groups. In contrast, in the present study, the FI of the spotted scat was

highly correlated with the Ulva content in their feed. The fish in the control group who

received no Ulva supplementation in their feed showed the lowest FI thus clearly

showing that Ulva sp. feed-supplementation affects FI in spotted scat. Generally, the FI
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stimulants found in seaweed are the non-essential amino acids, aspartic and glutamic

acid (Mabeau et al., 1992). In U. rigida , these amino acid were found to constitute the

highest content of any non-essential amino acid at levels of 1.15 and 0.91% for aspartic

and glutamic acid, respectively (Mahae et al., 2014). The positive effect on FI of Ulva-

supplemented feed will reduce the length of the weaning period of spotted scat which

will be of benefit in hatcheries because of the reduction on cost consequent upon the

lower use of Artemia.

The study of Abdel-Warith et al. (2016) found that the FI, WG, SGR and PER of

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9.59 ± 0.43 African catfish (Clarias gariepinus) fed with U. lactuca supplemented feed

for 55 weeks decreased and were negatively correlated with the seaweed content in the
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fish feed. In addition, The FCR was higher in fish that were fed with a high content of
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seaweed feed additive. However, the SGR, FCR and PER of the catfish fed with 55%
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Ulva-supplemented feed were not significantly different from those of fish in the non-

seaweed-supplemented control group and this was similar to the present study, in which
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the spotted scat fed the unsupplemented control diet and the 0 and 10% supplemented
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feed produced the best FCR performance. Therefore, supplementing the feed of spotted
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scat with Ulva in proportions of more than 10% may negatively affected feed quality.
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The FCE, parameter is calculated by comparing the weight gain with the weight of

feed available. Higher quality feed therefore produces a higher FCE than low quality

feed. In the present study, the highest FCE was achieved by the spotted scat fed with

5% Ulva-supplemented feed while the fish fed the 30% seaweed-supplemented feed and

the control group fish revealed the lowest FCE. This result is similar to that of Kut

Güroy et al. (2007) who studied Ulva rigida-feed supplementation in tilapia (O.

niloticus) and found that a lower content of Ulva in their feed (5–10%) was more

suitable for their culture. Meanwhile El-Tawil (2010) and Abdel-Warith et al. (2016)
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suggested that the most suitable level of Ulva sp. supplementation for tilapia

(Oreochromis sp.) and African catfish were 15% and 10 respectively. Finally, in spotted

scat, seaweed supplemented feed did not affected the SR of the fish which is similar to

the report of El-Tawil (2010) relating to tilapia.

This study clearly shows that Ulva sp. can stimulate the appetite of Scatophagus

argus while they develop from fry to juveniles, and it has the potential to be used as a

feedstock for spotted scat hatcheries. Further, its use as a feed supplement may reduce

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the weaning period. This seaweed can be added at levels of up to 25% to maximize

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WG, SGR, and FCE without significantly affecting SR as compared to fish fed an

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unsupplemented diet in the control group. However, the optimum level of dietary
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substitution of Ulva sp. would be at levels of 5–10% to achieve the best improvements

in growth performance and feed utilization.

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Conflict of Interest: The authors declare that they have no conflict of interest.

Ethical approval: All applicable national, and institutional guidelines for the care and
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use of animals were followed by the authors.

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Acknowledgements

We thank The Coastal Aquaculture Research and Development Regional Center 6,

Songkhla, Thailand for providing fish rearing facilities and Mr. Jarupong Makthunka

for rearing the fish throughout this study. We are also grateful to Mr. Michael Currie for

editing the English in the paper.

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Figure 1. Rate of feed intake in spotted scat fed the experimental diets for 10 weeks.
Data represent the mean ± SD. Significant differences at P < 0.05 (n = 20) between
groups are indicated by different letters over the bar graphs.
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Table 1 Nutritional composition of different raw materials used as the diet ingredients

Raw materials (% of dry weight)

Moisture Ash Crude protein Crude lipid Crude fiber

Fish meal 0.10±0.14 25.37±0.05 66.32±0.64 7.02±0.36 ND

Squid meal 0.48±0.06 8.20±0.07 49.62±0.09 13.31±0.52 ND

Soybean meal 1.00±0.37 7.18±0.06 50.04±0.21 0.99±0.05 7.83±0.30

Broken rice 2.15±0.09 0.07±0.03 8.85±0.01 0.30±0.02 0.74±0.06

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Wheat flour 0.47±0.12 0.19±0.04 13.59±0.08 0.90±0.08 0.60±0.10

Ulva rigida 0.36±0.17 24.68±0.18 pr

24.67±0.24 0.31±0.07 5.29±0.14
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Values are the mean ± SD of three replications, ND = not detected.
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Table 2 Ingredients and nutrient contents of the experimental diets

Ingredients (g/100 g) (% of dry weight)

1 2 3 4 5 6 7

Fish meal 22 21 19 19 16 16 16

Squid meal 13 12 13 12 15 11 12

Soybean meal 20 22 22 20 20 22 19

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Broken rice 11 10 12 11 10 9 10

Wheat flour 19 17 12 12 9 9 6

U. rigida 0 5 10
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Live fish oil 3 3 3 3 3 3 3
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Mineral mixa 2 2 2 2 2 2 2

Vitamin mixb
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2 2 2 2 2 2 2
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Rice husk 8 6 5 4 3 1 0

Nutrient contentsc (DM)

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Moisture 1.10± 1.07± 0.96± 1.08± 0.77± 0.69± 1.05±

0.14 0.05 0.18 0.19 0.18 0.09 0.11

Crude protein 39.61 39.72 39.33 39.52 39.38 39.73 39.63

±0.06 ±0.24 ±0.14 ±0.19 ±0.15 ±0.10 ±0.31

Crude lipid 6.13± 5.59± 5.66± 5.76± 5.41± 5.34± 5.37±

0.22 0.03 0.21 0.04 0.19 0.03 0.00

Crude fiber 2.10± 2.39± 2.87±. 2.53± 2.86± 2.85± 3.90±

0.05 0.02 0.00 0.02 0.02 0.14 0.01
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Ash 12.06 12.74 13.34 14.15 14.84 14.78 16.09

±0.03 ±0.03 ±0.17 ±0.06 ±0.06 ±0.05 ±0.14

Gross energyd 392.68 393.88 383.66 376.85 366.99 362.95 367.75

±2.38 ±0.08 ±1.42 ±0.51 ±0.48 ±0.70 ±0.37

a
Per kg of mineral mixture: iron, 12,000 mg; copper 12,000 mg; zinc, 15,000 mg;
manganese, 6,000 mg; iodine, 200 mg; selenium, 25 mg; magnesium 50,000 mg;
calcium, 100,000 mg; phosphorus, 80,000 mg.

b
Per kg of vitamin mixture: vitamin A, 600,000 IU; vitamin D3, 200,000 IU; vitamin E,

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6,000 IU; vitamin K, 1,200 mg; vitamin B1, 5,000 mg; vitamin B2, 6,000 IU; vitamin
B6, 5,000 mg; vitamin B12, 6 mg; niacin, 20,000 mg; pantothenic acid, 16,000 mg;

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folic acid, 1,000 mg; biotin, 200 mg; Endox Dry 20,000 mg.

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Nutrient content by proximate analysis.

d
kcal/100 g.
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Table 3 Amino acids in the experimental diets

Amino acid Experimental diets

(mg/100 mg) 1 2 3 4 5 6 7

Essential amino acids

Histidine 0.58± 0.55± 0.53± 0.53± 0.51± 0.49± 0.49±

0.03a 0.02ab 0.01bc 0.02bc 0.03cd 0.03cd 0.02d

Isoleucine 1.15± 1.14± 1.15± 1.14± 1.13± 1.10± 1.09±

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0.02a 0.01ab 0.01a 0.01a 0.04ab 0.04bc 0.02c

Leucine 2.13± 2.10± 2.10± 2.07± 2.09± 2.00± 2.02±

0.05a
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0.02a 0.03a 0.04abc 0.05ab 0.05c 0.03bc
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Lysine 1.55± 1.58± 1.59± 1.46± 1.53± 1.53± 1.64±
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0.08bc 0.05ab 0.02ab 0.06c 0.04bc 0.03bc 0.03a

Phenylalanine 1.19± 1.18± 1.16± 1.26± 1.22± 1.19± 1.18±

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0.07a 0.03a 0.02a 0.08a 0.06a 0.05a 0.02a

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Threonine 1.25± 1.26± 1.25± 1.26± 1.28± 1.23± 1.27±

0.03a 0.02a 0.02a 0.02a 0.03a 0.02a 0.02a

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Valine 1.31± 1.32± 1.32± 1.34± 1.34± 1.32± 1.32±

0.02a 0.02a 0.02a 0.00a 0.02a 0.03a 0.01a

Non essential amino acids

Alanine 1.73± 1.78± 1.80± 1.80± 1.85± 1.80± 1.90±

0.03d 0.03c 0.03c 0.03c 0.02b 0.01c 0.02a

Arginine 1.87± 1.83± 1.81± 1.86± 1.83± 1.79± 1.78±

0.09a 0.05a 0.03a 0.03a 0.07a 0.06a 0.01a

Aspartic acid 3.09± 3.19± 3.17± 3.12± 3.27± 3.12± 3.33±

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0.07c 0.13abc 0.05bc 0.05bc 0.08ab 0.09bc 0.09a

Glutamic acid 5.60± 5.63± 5.48± 5.21± 5.21± 4.95± 5.03±

0.09a 0.17a 0.06ab 0.05b 0.09b 0.12c 0.12bc

Glycine 1.71± 1.73± 1.70± 1.75± 1.71± 1.62± 1.66±

0.06ab 0.04ab 0.03ab 0.03a 0.06ab 0.04c 0.03bc

Proline 1.53± 1.52± 1.48±. 1.47± 1.46± 1.41± 1.40±

0.04a 0.01ab 0.03ab 0.03ab 0.03bc 0.05c 0.02c

Serine 1.37± 1.39± 1.36± 1.39± 1.39± 1.36± 1.39±

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0.03a 0.03a 0.03a 0.02a 0.03a 0.05a 0.02a

Tyrosine 0.67± 0.66± 0.65±. 0.70± 0.68± 0.66± 0.66±

0.03a
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0.02a 0.02a 0.04a 0.03a 0.04a 0.02a
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Total 26.72 26.87 26.54 26.36 26.51 25.59 26.16
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±0.42a ±0.28a ±0.14a ±0.12a ±0.19a ±0.56c ±0.21

b b b b
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Values are the mean ± SD of three replications.

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Mean within each row not sharing a common superscript are significantly different (P <
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0.05).
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Table 4 Growth performance and survival rate of spotted scat fed the experimental diets
for 10 weeks
Seaweed Weight gain Specific growth rate (%/ Survival

(%) (%) fish/day) (%)

0 434.97±23.62cd 2.39±0.06cd 98.33±2.89a

5 593.91±18.13a 2.77±0.04a 98.33±2.89a

10 536.85±38.85ab 2.64±0.09ab 96.67±5.77a

15 484.67±23.88bc 2.52±0.06bc 98.33±2.89a

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20 474.19±19.46bc 2.50±0.05bc 98.33±2.89a

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25 445.88±96.96cd 2.41±0.26cd 100.00±0.00a

30 369.25±22.34d pr 2.21±0.07d 100.00±0.00a

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Values are the mean ± SD of three replications.
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Mean within each column not sharing a common superscript are significantly different
(P < 0.05).
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Table 5 Feed utilization of spotted scat fed the experimental diets for 10 weeks

Seaweed Feed Feed Protein Lipid Energy

(%) conversion conversion efficiency efficiency efficiency
rate efficiency (%) ratio ratio ratio (g/kcal)

0 2.25±0.12c 9.77±1.00b 1.12±0.06a 7.26±0.55ab 0.113±0.006a

5 2.26±0.11c 12.20±1.73a 1.12±0.05a 7.92±0.37a 0.113±0.006a

10 2.52±0.18bc 10.82±1.45ab 1.07±0.10ab 7.05±0.73ab 0.104±0.008ab

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15 2.64±0.13ab 10.72±1.12ab 0.96±0.05bc 6.59±0.29b 0.101±0.005ab

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20 2.61±0.16ab 11.29±1.14ab 0.97±0.06bc 7.09±0.36ab 0.105±0.006ab

25 2.73±0.25ab 9.87±2.71ab
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0.93±0.08c 6.88±0.59b 0.101±0.009ab
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30 2.90±0.11a 8.79±0.91b 0.87±0.04c 6.42±0.25b 0.094±0.004b
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Values are the mean ± SD of three replications.

Mean within each column not sharing a common superscript are significantly different
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(P < 0.05).
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Highlights
 U. rigida can stimulate the appetite of Scatophagus argus while they develop from fry
to juveniles.

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