Aquaculture 501 (2019) 368–373

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Aquaculture
journal homepage: www.elsevier.com/locate/aquaculture

Effects of dietary protein and lipid levels on growth, body composition and T
nutrient utilization of Channa striata
Katheline Huaa,
⁎,1
, Wolfgang Koppeb,2, Ramon Fontanillasb
a
Faculty of Life Sciences, Humboldt-Universität zu Berlin, Invalidenstraße 42, 10115 Berlin, Germany
b
Skretting Aquaculture Research Centre, Sjøhagen 3, 4016 Stavanger, Norway

ARTICLE INFO ABSTRACT

Keywords: Very limited information is available on nutrient requirements of Channa striata, particularly during the grow-
Protein requirement out stage. A study was carried out to study the requirement and utilization of protein and lipid by Channa striata
Lipid requirement growing from 31 g to ~ 200 g. Nine experimental diets with graded levels of dietary protein and lipid were
Protein-sparing effect formulated using practical ingredients. Increasing dietary crude protein content from 43.4% to 55.6% at 11%
Nutrient utilization
dietary lipid content resulted in increases of weight gain, amount of feed consumed, specific growth rate, and
thermal-unit growth coefficient. The crude protein requirement of Channa striata was estimated by the saturation
kinetics model to be 51.6% dry matter (DM) for growth and 52.1% DM for protein deposition. On the digestible
protein basis, the requirement of Channa striata was estimated to be 47.2% DM for growth and 47.8% DM for
protein deposition. Increasing lipid content in diets from 7.5% to 23.1% in this study resulted in linear decreases
of fish growth and feed intake. Apparently Channa striata cannot tolerate high dietary lipid levels; lipid levels
of > 19% DM could negatively affect growth performance. It was also revealed that dietary lipids did not have a
protein-sparing effect in Channa striata. Results from this study not only contribute to a better understanding of
the nutrient requirement and utilization of Channa striata, but also would be helpful for the development of cost-
effective sustainable practical feed formulations for this fish species.

1. Introduction
The striped snakehead, Channa striata (Bloch, 1793), is a high-value
food fish species that is widely cultured in Southeast Asia
(Paripatananont, 2002). It is also used in traditional medicine due to its
pharmaceutical properties (Paripatananont, 2002). They are generally
farmed in intensive culture systems and harvested when they reach a
market size of 600 to 700 g (Hung and Huy, 2007). It is an air-breathing
species that is very hardy and tolerant to low dissolved oxygen levels
and thus is often cultured in areas unsuitable for other fish species
(Samantaray and Mohanty, 1997). Channa striata is a carnivorous spe-
cies. In practice, they are traditionally fed farm-made feeds that contain
small fresh fish as main feed ingredients (Hung and Huy, 2007). For-
mulated feeds of Channa striata generally contain high levels of in-
gredients from animal origin, particularly fish meals. A high level of
fish meal in fish feeds not only could potentially put an ecological
pressure on wild fish stocks, but also could result in an elevated feed
cost since fish meal is an expensive ingredient. Feed cost in intensive
Channa striata culture systems can reach 70 to 80% of total production
cost (Hung and Huy, 2007). Numerous studies have been conducted to
investigate the effect of replacing fish meal by alternative protein
sources in various fish species, including salmonids, seabass, seabream,
etc. (NRC, 2011). So far only a few studies have been carried out to
study the effect of replacing fish meal in Channa striata feeds using
practical ingredients, including poultry by-product meal (Abdul-Halim
et al., 2014), soybean meal (Hien et al., 2015) and soy protein con-
centrate (Hien et al., 2017).
This, in part, is due to our limited knowledge on Channa striata
nutrition at present. Only a handful of studies have been published on
the requirement and utilization of protein and lipid by Channa striata.
The protein requirement of Channa striata has been reported to be 55%
crude protein or 50% digestible protein (Mohanty and Samantaray,
1996). The optimal lipid level was found to be below 11.5% (Aliyu-
Paiko et al., 2010) and 13% (Samantaray and Mohanty, 1997).

Corresponding author.
⁎

E-mail address: katheline.hua@jcu.edu.au (K. Hua).

1
Present address: Tropical Futures Institute, James Cook University Singapore, 149 Sims Drive, 387380 Singapore; Centre for Sustainable Tropical Fisheries and
Aquaculture, James Cook University, 1 James Cook Drive, Townsville QLD 4811, Australia.
2
Present address: Simplyfish AS, Baldersgate 12, 4011 Stavanger, Norway.

https://doi.org/10.1016/j.aquaculture.2018.11.054
Received 13 September 2018; Received in revised form 28 November 2018; Accepted 28 November 2018
Available online 29 November 2018
0044-8486/ © 2018 Elsevier B.V. All rights reserved.
K. Hua et al. Aquaculture 501 (2019) 368–373

Table 1
Formulation of the experimental diets.
Ingredients (%) Diets

43/11 46/11 49/11 52/11 55/11 52/7 52/15 52/19 52/23

Wheat 45.82 40.34 35.73 29.95 24.71 33.21 24.95 19.93 14.58
Soybean meal 9.35 10.50 11.50 12.50 13.50 12.50 12.50 12.50 12.50
Wheat gluten 2.00 2.00 2.00 2.00 2.00 1.49 2.98 3.89 6.75
Fish meal 18.30 20.05 21.50 23.79 25.45 23.70 23.70 23.70 23.70
Poultry meal 11.00 12.50 14.00 16.00 18.00 16.00 16.00 16.00 16.00
Feather meal 4.00 5.00 5.50 6.00 6.50 6.00 6.00 6.00 5.40
Hemoglobin 4.00 4.50 5.00 5.50 6.00 5.50 5.50 5.50 4.50
Rapeseed oil 1.30 1.16 1.05 0.88 0.74 0.00 2.23 3.59 4.94
Fish oil 2.63 2.35 2.12 1.78 1.50 0.00 4.54 7.29 10.03
Yttrium oxide 0.10 0.10 0.10 0.10 0.10 0.10 0.10 0.10 0.10
Vitamin and mineral premix1 1.50 1.50 1.50 1.50 1.50 1.50 1.50 1.50 1.50
Total 100 100 100 100 100 100 100 100 100

1
Skretting standard vitamin and minerals premix, according to requirement data from NRC (2011). Trouw Nutrition, the Netherlands.

However, these studies mainly focused on the nutrition of fry or juve-
nile fish at body weights below 20 g. It is during the grow-out stage that
fish consume the majority of feeds throughout the production cycle,
which has a significant bearing on the economic and ecological sus-
tainability of Channa striata culture. Therefore, it is essential to study
the nutrient requirement and utilization and develop sustainable feed
formulations for Channa striata during this crucial life stage.
The objectives of this study were to study the requirement and
utilization of protein and lipid by Channa striata during the grow-out
stage.
2. Materials and methods
2.1. Experimental diets
Nine experimental diets were formulated with practical ingredients
(Table 1). All experimental diets were based on fish meal (ranging from
18.3% to 25.45% of diets) combined with ingredients from plant
sources (soybean meal and wheat) and animal by-products (poultry
meal, feather meal and hemoglobin). Five diets were formulated to
contain graded levels of crude protein (CP) at 43, 46, 49, 52, and 55%
of dry matter (DM) while maintaining the lipid content at 11% (DM).
Another four diets were formulated to contain graded levels of lipid at
7, 15, and 19 and 23% (DM) while maintaining the crude protein
content at 52% (DM). Combined with one of the diets from the protein
series, there were five graded levels of dietary lipid at the crude protein
content of 52%. The analyzed chemical composition of the experi-
mental diets is presented in Table 2. The experimental diets were pro-
duced by extrusion at the Skretting Aquaculture Research Centre
Technology Plant (Stavanger, Norway).
diets. During the growth trial, digestibility of the experimental diets
was determined using a modified Guelph system (Cho et al., 1982).
Fecal samples were collected according to Cho et al. (1982) during the
last 6 weeks of the trial.
2.3. Sampling and chemical analyses
Fish carcass samples were taken at the beginning and end of the
growth trial. Fish were euthanized by an overdose of MS222 (Ethyl 3-
aminobenzoate). The fish were pooled by tanks, ground into homo-
geneous slurry, freeze-dried, reground and stored at −20 °C until
analyzed. Diet, ingredients and carcass samples were analyzed for dry
matter (DM) and ash according to AOAC (1995), crude protein (% N x
6.25) by the Dumas combustion principle (LECO Corporation, St. Jo-
seph, MI, USA). Total lipid content was measured by Maran Ultra NMR
nuclear magnetic resonance (Resonance Instruments Ltd., Witney, UK).
Gross energy content of samples was measured by Masterlab (Boxmeer,
the Netherlands). Contents of yttrium, calcium, phosphorus, potassium
and magnesium were analyzed by inductively coupled plasma optical
emission spectrometry (ICP-OES). Content of amino acids were de-
termined by ion-exchange chromatography (Commission Directive,
1998).
2.4. Calculations
Live weight gain, growth rate, and feed efficiency ratio (live weight
gain: feed dry weight) were calculated. Growth rate was calculated
using specific growth rate (SGR) and thermal-unit growth coefficient
(TGC) (Iwama and Tautz, 1981) according to the following equations:
SGR = [ln FBW ln IBW ]/d × 100

TGC = [FBW (1/3) IBW (1/3) ]/[ T × d ) ] × 100

2.2. Fish, experimental conditions, and feeding
The trial was conducted at Humboldt-Universität zu Berlin in a re-
circulation system for 83 days. Channa striata were obtained from a
commercial hatchery (Trai Ca Loc Giong 2 Lam, Vietnam). Triplicate
groups of fish with an average body weight (BW) of 31.3 ± 0.2 g were
randomly allocated to each of the nine experimental diets. The re-
circulation system consisted of 30 fiberglass tanks with a rearing vo-
lume of 200 l. Water temperature was maintained at 27.7 °C. A photo-
period regime of 12 h light and 12 h was maintained by artificial
lighting. The fish were treated in accordance with the German law on
animal experiments.
Fish were hand-fed the experimental diets to excess twice a day
during weekdays and once a day during weekends. Excess feeds were
collected and oven-dried after each meal and were used to calculate the
actual feed intake by taking into account the dry matter content of the
where FBW = final body weight (g), IBW = initial body weight (g),
T = water temperature (°C), and d = the number of days.
The apparent digestibility coefficients (ADC) for the nutrients and
energy of the diet were calculated as follows (Cho et al., 1982):
ADC = 1 (F/D Di/Fi)
where: D = % nutrient (or kJ/g gross energy) of diet; F = % nutrient
(or kJ/g gross energy) of feces; Di = % digestion indicator of diet;
Fi = % digestion indicator of feces.
2.5. Statistical analysis
Data were analyzed as a complete random design using the general
linear model (GLM) of the SAS/STAT software package version 9.3 (SAS
Institute, Cary, NC, USA). Means of dependent variables were compared

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K. Hua et al. Aquaculture 501 (2019) 368–373

Table 2
Analyzed chemical composition of experimental diets (DM basis).
Diets

43/11 46/11 49/11 52/11 55/11 52/7 52/15 52/19 52/23

Crude protein (%) 43.4 47.2 49.2 51.9 55.6 53.1 52.5 52.0 52.2
Crude lipid (%) 10.7 10.6 10.3 10.4 10.5 7.5 14.6 19.0 23.1
Ash (%) 6.6 7.2 7.4 8.1 8.6 8.1 7.8 7.7 7.7
Gross energy (MJ/kg) 21.0 21.1 21.0 21.1 21.3 20.7 22.2 23.1 24.2

AAs
Arginine (%) 2.5 2.7 2.9 3.1 3.4 3.2 3.2 3.1 3.1
Histidine (%) 1.1 1.2 1.2 1.3 1.5 1.4 1.4 1.3 1.3
Isoleucine (%) 1.5 1.6 1.7 1.9 2.0 1.9 1.9 1.9 1.9
Leucine (%) 3.4 3.6 3.8 4.1 4.4 4.2 4.1 4.1 4.0
Lysine (%) 2.6 2.5 3.0 2.9 3.1 2.9 2.9 2.9 2.8
Methionine (%) 0.9 0.9 0.9 0.9 0.9 0.9 0.9 0.9 0.9
Cystine (%) 0.7 0.8 0.8 0.9 0.9 0.9 0.9 0.9 0.9
Phenylalanine (%) 1.9 2.1 2.2 2.2 2.4 2.3 2.3 2.3 2.3
Tyrosine (%) 1.0 1.1 1.1 1.2 1.2 1.2 1.2 1.2 1.2
Threonine (%) 1.6 1.8 1.8 2.0 2.1 2.0 2.0 2.0 1.9
Valine (%) 2.1 2.3 2.5 2.7 2.9 2.7 2.7 2.7 2.6
Alanine (%) 2.3 2.5 2.7 2.9 3.1 3.0 2.9 2.9 2.8
Aspartic acid (%) 3.5 3.8 4.0 4.3 4.6 4.4 4.3 4.3 4.2
Glutamic acid (%) 7.0 7.4 7.6 7.7 8.1 7.9 7.9 7.9 8.3
Glycine (%) 2.5 2.8 3.0 3.2 3.5 3.3 3.3 3.2 3.2
Proline (%) 2.7 3.0 3.1 3.2 3.4 3.3 3.3 3.3 3.4
Serine (%) 2.2 2.5 2.6 2.8 2.9 2.9 2.8 2.9 2.8
Sum of AAs (%) 39.6 42.6 44.9 47.2 50.4 48.6 47.9 47.6 47.5

using Tukey's honestly significant difference (HSD) test, significance Table 3

was considered when p < .05. Apparent digestibility coefficients (ADCs) of the experimental diets containing
Saturation kinetics model (Mercer, 1982) was used to estimate increasing levels of dietary protein.
protein requirement of Channa striata from the body weight gain and ADC (%) Diets SEM
the protein gain of fish fed graded protein levels. The saturation kinetics
model is described as follows: 43/11 46/11 49/11 52/11 55/11

d (k 0.5 )n + ax n Crude protein 89.8b 91.4ab 91.4ab 92.0ab 93.0a 0.59

y= Crude lipid 88.5 89.8 90.5 91.6 93.5 1.17
(k 0.5 ) n + x n
Gross energy 82.4c 84.9bc 85.3abc 86.1ab 88.4a 0.87
where, Arginine 93.1b 94.4b 94.4b 95.4a 95.9a 0.39
Histidine 92.3b 94.6ab 94.4a 95.2a 95.3a 0.46
y = body weight gain (g), or protein gain (g). Isoleucine 88.5b 90.9ab 90.5ab 91.9a 92.6a 0.62
Leucine 91.4 93.1 92.8 93.7 94.3 0.63
x = crude or digestible protein content (% DM).
Lysine 94.1 95.0 95.5 95.6 96.1 0.43
a = plateau of the curve (upper asymptote). Methionine 92.5 93.5 92.5 93.1 93.3 0.57
d = intercept on y-axis. Cystine 83.4b 85.4ab 85.5ab 87.1a 88.3a 0.66
k0.5 = concentration for ½ of (a + d). Phenylalanine 90.7b 92.8ab 92.2ab 93.1ab 93.8a 0.50
n = apparent kinetic order. Tyrosine 91.3b 93.0ab 93.3ab 93.6ab 95.0a 0.53
Threonine 87.8b 90.5a 90.4a 91.7a 92.5a 0.53
Valine 88.5b 91.2ab 90.8ab 92.1a 93.0a 0.67
The analyses were carried out with GraphPad Prism software ver- Alanine 87.9b 91.3ab 90.8a 92.2a 93.5a 0.65
sion 7 (GraphPad Software, La Jolla, CA, USA). Aspartic acid 90.7b 93.1ab 92.8ab 93.6a 94.0a 0.56
Glutamic acid 94.9b 96.0ab 95.7ab 96.2ab 96.6a 0.30
Glycine 88.0b 91.2ab 91.4a 92.8a 93.3a 0.69
3. Results Proline 91.3b 92.9ab 93.1ab 94.0a 94.7a 0.58
Serine 88.5a 91.1ab 91.0ab 92.5a 93.6a 0.56
The ADCs of the experimental diets are presented in Tables 3 and 4. Sum of AAs 91.1b 93.0ab 92.9ab 93.7a 94.4a 0.50
ADCs of protein and energy were significantly and positively affected
by increasing levels of dietary protein. However, dietary protein level Values in the same row not sharing the same letter are significantly different;
SEM = Pooled standard error of the mean.
did not affect the digestibility of dietary lipid. Dietary protein level
significantly and positively affected digestibility of all amino acids ex-
and 55% protein. The increase in dietary crude protein content resulted
cept for leucine, lysine and methionine. Increasing dietary lipid levels
in a linear increase in feed efficiency from 0.90 to 1.20. Feed intake was
significantly improved lipid and energy digestibility, but had no effect
not significantly affected by dietary crude protein levels although there
on protein digestibility. Digestibility of most of the amino acids were
was a trend of increase in feed intake with increasing dietary crude
significantly affected by dietary lipid levels except for methionine,
protein levels. In contrast, increasing dietary lipid levels significantly
alanine, aspartic acid, glycine and proline.
depressed feed intake of fish, as well as final body weight, body weight
Tables 5 and 6 present fish growth performance and feed efficiency.
gain, SGR, TGC. The depressing effects were significant when the lipid
Body weight gain, SGR and TGC were positively affected by dietary
level reached 19%, but there were no statistical differences among the
protein levels. Diets containing 52 and 55% crude protein showed
dietary treatments of 7, 11, and 15% of dietary lipid levels. Dietary lipid
significantly higher body weight gain, SGR and TGC compared to diets
levels did not have a significant effect on feed efficiency when the
with 43% crude protein. However, there were no significant differences
dietary protein content was held constant.
in body weight gain, SGR and TGC among the diets containing 49, 52

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K. Hua et al. Aquaculture 501 (2019) 368–373

Table 4 carcass lipid content from 5.7 to 9.9% and energy content from 6.8 to
Apparent digestibility coefficients (ADCs) of the experimental diets containing 8.3 kJ/g. Ash content and mineral components (Ca, P, K, Mg) were not
increasing levels of dietary lipid. affected by dietary treatments.
ADC (%) 52/7 52/11 52/15 52/19 52/23 SEM Tables 9 and 10 present the nutrient retention efficiency. Nitrogen
retention efficiency was high for fish fed all experimental diets, ranging
Crude Protein 92.8 92.0 92.6 93.0 93.5 0.39 from 40 to 44% when expressed as apparent nitrogen retention effi-
Crude lipid 90.9c 91.6bc 94.1ab 95.8a 95.5a 0.67
ciency, and from 45 to 48% when expressed as net nitrogen retention
Gross energy 87.0cd 86.1d 87.8bc 89.2ab 89.9a 0.37
efficiency. Neither dietary protein nor lipid levels affected nitrogen
Arginine 95.3b 95.4ab 95.5ab 96.1a 96.0ab 0.15
retention efficiency. Dietary protein levels significantly affected energy
Histidine 95.0b 95.2ab 94.5ab 95.4a 95.4a 0.16
Isoleucine 91.8b 91.9ab 91.8ab 92.9a 93.0a 0.24 retention efficiency. Apparent energy retention efficiency increased
Leucine 93.6b 93.7b 93.6b 94.7a 94.9a 0.17 from 31.3 to 41.8%, whereas net energy retention efficiency increase
Lysine 95.5c 95.6bc 95.7abc 96.1ab 96.2a 0.12 from 38.0 to 47.3% with the increase of dietary protein content.
Methionine 93.2 93.1 93.1 94.0 93.5 0.22 However, dietary lipid levels did not affect apparent or net energy re-
Cystine 87.7b 87.1b 89.0a 90.1a 89.8a 0.25
Phenylalanine 93.2bc 93.1c 93.1c 94.1ab 94.5a 0.21
tention efficiency.
Tyrosine 94.0ab 93.6b 94.6a 94.8a 94.6a 0.21 On the crude protein basis, the requirement was estimated to be
Threonine 92.0ab 91.7ab 91.3b 92.5a 92.3ab 0.19 51.6% DM for body weight gain, and 52.1% DM for protein gain with
Valine 92.4abc 92.1c 92.2bc 93.2ab 93.3a 0.22 the saturation kinetics model. The protein requirement was estimated
Alanine 92.6 92.2 92.3 93.6 93.3 0.29
to be 47.2% DM digestible protein by the saturation kinetics model
Aspartic acid 93.8 93.6 93.4 94.0 93.8 0.14
Glutamic acid 96.2c 96.2c 96.4bc 96.8ab 96.9a 0.09 based on body weight gain (Fig. 1), 47.8% DM digestible protein by the
Glycine 92.9 92.8 92.6 93.2 93.1 0.30 saturation kinetics model based on protein gain (Fig. 2).
Proline 94.1 94.0 94.0 94.8 95.0 0.25
Serine 92.5b 92.5b 93.0ab 94.0a 94.0a 0.20 4. Discussion
Sum of AAs 93.9b 93.7b 93.8b 94.6a 94.6a 0.14

Values in the same row not sharing the same letter are significantly different; This study examined the effect of increasing dietary crude protein
SEM = Pooled standard error of the mean. content at constant dietary lipid content and the effect of increasing
dietary lipid content at constant dietary protein content in Channa
Table 5 striata growing from 31 to ~ 200 g. This was achieved through nine
Growth performance and feed efficiency of Channa striata1 fed increasing levels experimental diets with five graded levels of dietary protein and lipid
of dietary protein for 12 weeks. each.
In this study, increasing dietary protein content from 43.4% to
Diet FBW BWG Feed FE SGR TGC
55.6% at 11% dietary lipid content resulted in increases of weight gain,
g/fish g/fish g/fish gain: feed %/day amount of feed consumed, SGR and TGC. Fish fed diets 43/11 and 46/
c c d
11 had significant lower body weight and growth rate than those fed
43/11 147.7 116.7 129.4 0.90 1.88c 0.093c
diets 49/11, 52/11 and 55/11. It appears that the crude protein re-
46/11 160.6bc 130.2bc 127.5 1.02c 2.00bc 0.101bc
49/11 183.9abc 152.3abc 142.6 1.07bc 2.12abc 0.110abc quirement of Channa striata is in the range between 49.2 and 55.6%.
52/11 200.1ab 168.6ab 150.6 1.12b 2.23ab 0.117ab This corresponds to a digestible protein requirement between 45.0 and
55/11 209.7a 177.9a 148.7 1.20a 2.27a 0.120a 51.7%. Net nitrogen retention efficiency of fish in this study remained
SEM 9.1 8.8 7.1 0.02 0.05 0.004 high and unaffected by dietary protein content; in the meantime the
crude protein content in whole fish body was also unaffected by in-
1
Initial live weight of fish = 31.3 g; values in the same column not sharing creasing dietary protein levels. Apparently Channa striata was able to
the same letter are significantly different; SEM = Pooled standard error of the effectively utilize dietary proteins at all protein levels tested in this
mean.
study.
The protein requirement was further estimated in the present study
Table 6 using the saturation kinetics model. Estimates of protein requirement
Growth performance and feed efficiency of Channa striata1 fed increasing levels
differed slightly according to the response criteria (body weight gain
of dietary lipid for 12 weeks.
versus protein deposition). The estimates based on body weight gain
Diet FBW BWG Feed FE SGR TGC tended to be lower than the estimates based on protein gain. Based on
the saturation kinetics model, the crude protein requirement of Channa
g/fish g/fish g/fish gain: feed %/day
striata was estimated to be 51.6% DM for growth and 52.1% DM for
52/7 208.0 a
177.2 a
149.0 a
1.19 2.30a 0.121a protein deposition, and the digestible protein requirement to be 47.2%
52/11 200.1ab 168.6ab 150.6a 1.12 2.23ab 0.117ab DM for growth and 47.8% DM for protein deposition.
52/15 183.5abc 151.8abc 130.0ab 1.17 2.11abc 0.109abc The estimates of the protein requirement are in line with literature
52/19 156.2c 125.1c 107.2b 1.17 1.94c 0.097c
52/23 163.3bc 131.7bc 111.4b 1.18 1.98bc 0.100bc
reports for Channa striata and other species in the same family. For
Channa striata fry, the protein requirement was observed to be 55%
SEM 9.1 9.0 7.1 0.03 0.06 0.004
crude protein or 50% digestible protein on as is basis at the dietary lipid
1
Initial live weight of fish = 31.3 g; values in the same column not sharing level of 12% (Mohanty and Samantaray, 1996).Wee and Tacon (1982)
the same letter are significantly different; SEM = Pooled standard error of the estimated that the requirement was 52% crude protein for Channa mi-
mean. cropeltes. Based on weight gain, crude protein requirements of 50.5 and
47.5% DM were observed for fingerlings of hybrid snakehead (Channa
Carcass composition of the fish after 12 weeks is presented in Tables maculata ♀ × Channa argus ♂) at lipid levels of 6.5% and 12%, re-
7 and 8. Dietary protein levels did not have significant effects on the spectively (Zhang et al., 2017). Sagada et al. (2017) concluded that the
nutrient composition of the fish, although there was a trend of increase optimal protein level for juvenile northern snakehead (Channa argus)
in body protein concentration. On the other hand, dietary lipids levels was 48% (as is basis) when diets contained either 12% or 15% lipids.
significantly affected the contents of moisture, lipid and energy in fish On the other hand, a lower protein requirement of 43% was reported
body. Increasing lipid levels in diets resulted in a linear increase in for snakehead fry (unspecified Channa species) (Paripatananont, 2002).
The crude protein requirement was determined to be 45% DM for

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K. Hua et al. Aquaculture 501 (2019) 368–373

Table 7
Chemical composition of whole carcass of Channa striata1 fed increasing levels of dietary protein for 12 weeks.
Diet Moisture (%) CP (%) Lipid (%) Energy (KJ/g) Ash (%) Ca (%) P (%) K (%) Mg (%)

43/11 69.1 19.2 6.6 7.1 4.6 1.34 0.83 0.33 0.05
46/11 69.3 19.3 6.7 7.1 4.1 1.39 0.83 0.32 0.05
49/11 69.6 19.3 6.6 7.1 4.4 1.28 0.78 0.32 0.05
52/11 69.0 19.7 7.0 7.1 4.8 1.22 0.76 0.32 0.05
55/11 69.1 19.8 6.8 7.3 4.3 1.23 0.76 0.33 0.04

SEM 0.36 0.22 0.16 0.12 0.20 0.04 0.02 0.005 0.001

1
Initial body composition of fish: moisture 71.4%, CP 18.3%, lipid 5.3%, energy 6.5 kJ/g, ash 4.6%, Ca 1.32%, P 0.8%, K 0.3%, Mg 0.05%; values in the same
column not sharing the same letter are significantly different; SEM = Pooled standard error of the mean.

Table 8
Chemical composition of whole carcass of Channa striata1 fed increasing levels of dietary lipid for 12 weeks.
Diet Moisture (%) CP (%) Lipid (%) Energy (KJ/g) Ash (%) Ca (%) P (%) K (%) Mg (%)

a d c
52/7 70.2 19.6 5.7 6.8 4.2 1.24 0.77 0.33 0.05
52/11 69.0ab 19.7 7.0cd 7.1bc 4.8 1.22 0.76 0.32 0.05
52/15 68.6ab 19.4 8.1bc 7.6abc 4.7 1.14 0.71 0.32 0.04
52/19 67.7ab 19.2 8.8ab 7.9ab 4.6 1.26 0.77 0.32 0.04
52/23 66.5b 19.3 9.9a 8.3a 4.6 1.25 0.76 0.32 0.04

SEM 0.68 0.33 0.31 0.24 0.31 0.04 0.02 0.01 0.001

1
Initial body composition of fish: moisture 71.4%, CP 18.3%, lipid 5.3%, energy 6.5 kJ/g, ash 4.6%, Ca 1.32%, P 0.8%, K 0.3%, Mg 0.05%; Values in the same
column not sharing the same letter are significantly different; SEM = Pooled standard error of the mean.

Table 9
Nutrient retention efficiency of Channa striata fed increasing levels of dietary
protein for 12 weeks.
Diet Protein gain NRE NRE Energy gain ERE ERE

(g) (% NI) (%DNI) (kJ) (% EI) (%DEI)

b c c
43/11 22.8 40.4 45.0 851.2 31.3 38.0c
46/11 25.3b 42.1 46.1 938.7bc 34.8bc 41.0bc
49/11 29.7ab 42.3 46.3 1103.1abc 36.7b 43.0ab
52/11 33.6a 43.0 46.4 1218.5ab 38.5ab 44.7ab
55/11 35.7a 43.2 46.8 1327.3a 41.8a 47.3a

SEM 2.40 0.88 0.88 98.61 1.21 1.31

Values in the same column not sharing the same letter are significantly dif-
ferent; NRE = nitrogen retention efficiency, NI = nitrogen intake,
DNI = digestible nitrogen intake; ERE = energy retention efficiency,
EI = energy intake, DEI = digestible energy intake; SEM = Pooled standard
error of the mean.
Fig. 1. Live weight gain (g/fish ± SE) in response to dietary digestible protein
Table 10 levels after 12 weeks. The saturation kinetics model estimated the digestible
Nutrient retention efficiency of Channa striata fed increasing levels of dietary protein requirement to be 47.2% of dry matter. Requirement was established at
lipid for 12 weeks. achieving 95% of maximum performance.
Diet Protein gain NRE NRE Energy gain ERE ERE
content or the efficiency of protein utilization was not influenced by
(g) (% NI) (%DNI) (kJ) (% EI) (%DEI)
dietary lipid content in the present study. Increasing dietary lipid re-
52/7 35.2a 44.4 47.9 1214.0 39.3 45.2 sulted in a significant increase of body lipid content, indicating dietary
52/11 33.6ab 43.2 46.4 1218.5 38.5 44.7 lipid was directed towards body lipid deposition rather than towards
52/15 29.9abc 43.7 47.2 1183.3 40.8 46.4 protein-sparing. A lack of protein-sparing effect was also observed in
52/19 24.3c 43.8 47.1 1022.8 41.5 46.6
hybrid snakehead (Channa maculata ♀ × Channa argus ♂) (Zhang
52/23 25.7bc 44.2 47.3 1154.3 42.9 47.7
et al., 2017). In contrast, Sagada et al. (2017) observed a protein-
SEM 2.63 1.58 1.67 105.9 2.37 2.66 sparing effect in juvenile northern snakehead (Channa argus) when the
dietary lipid content increased from 9% to 12%.
Values in the same column not sharing the same letter are significantly dif-
ferent; NRE = nitrogen retention efficiency, NI = nitrogen intake, In addition to the lack of protein-sparing effect, the significant de-
DNI = digestible nitrogen intake; ERE = energy retention efficiency, crease in growth performance of fish fed high levels of dietary lipids in
EI = energy intake, DEI = digestible energy intake; SEM = Pooled standard the present study suggested that Channa striata cannot tolerate high
error of the mean. dietary lipid levels. Lipid levels of > 19% DM could be detrimental to
fish growth performance. This is in line with the observations from
Channa punctatus fingerlings (Zehra and Khan, 2012). several studies with Channa striata.Aliyu-Paiko et al. (2010) observed
Increasing dietary lipid from 7.5% to 23.1% in this study resulted in linearly depressed growth performance of Channa striata fingerlings fed
linear decreases of fish growth and feed intake. The body protein three lipids levels (6.5, 9 and 11.5%), and suggested that dietary lipid

372
K. Hua et al.
Fig. 2. Protein gain (g/fish ± SE) in response to dietary digestible protein le-
vels after 12 weeks. The saturation kinetics model estimated the digestible
protein requirement to be 47.8% of dry matter. Requirement was established at
achieving 95% of maximum performance.
should be lower than 11.5%. Samantaray and Mohanty (1997) fed diets
containing 9, 13, and 17% of lipids to Channa striata fingerlings and
concluded that 13% could be effectively utilized by fish. In contrast,
juvenile hybrid snakehead (Channa argus x Channa maculata) seemed to
be able to effectively utilize dietary lipids as their growth performance
was significantly improved when dietary lipid content increased from
5.8 to 17.3% at a dietary crude protein level of 42% (Zhao et al., 2016).
The present study observed significant effects of dietary protein
level on ADCs of protein, most of the amino acids, and energy. In ad-
dition, increasing dietary lipid levels significantly affected ADCs of
lipid, most of the amino acids, and energy. Since the protein levels and
lipid levels were achieved with different protein and lipid sources in the
present study, the significant effects cannot be attributed to protein
levels or lipid levels alone and the results should be taken with caution.
Nonetheless, the ADC values indicated that the dietary nutrients were
highly digestible in this study. Very little information exists for Channa
striata in literature regarding the digestibility of nutrients. To the best
knowledge of the authors, there has been no report on digestibility of
amino acids for Channa striata. The results from the present study
suggested that Channa striata could effectively digest and utilize nu-
trients from diets containing a combination of practical ingredients.
5. Conclusion
The present study suggested that diets containing protein and lipid
contents of 49/11, 52/11, 55/11, 52/7 and 52/15 could support good
growth performance of grow-out Channa striata. Channa striata required
crude protein and digestible protein at 51.6% DM and 47.2% for
growth, respectively. Based on the response criterion of the optimal
protein deposition, the crude protein and digestible protein require-
ments were 52.1% DM and 47.8% DM, respectively. Channa striata does
373
Aquaculture 501 (2019) 368–373
not seem to tolerate > 19% of lipids in their diets and apparently
dietary lipids do not spare dietary proteins from catabolism.
Acknowledgements
Financial support of this study was provided by Skretting
Aquaculture Research Centre, Norway. Technical assistance from the
staff at the Humboldt-Universität zu Berlin is thankfully acknowledged.
References
Abdul-Halim, H.H., Aliyu-Paiko, M., Hashim, R., 2014. Partial replacement of fish meal
with poultry by-product meal in diets for snakehead, Channa striata (Bloch, 1793),
fingerlings. J. World Aquacult. Soc. 45, 233–241.
Aliyu-Paiko, M., Hashim, R., Shu-Chien, A.C., 2010. Influence of dietary lipid/protein
ratio on survival, growth, body indices and digestive lipase activity in snakehead
(Channa striata, Bloch 1793) fry reared in re-circulating water system. Aquac. Nutr.
16, 466–474.
AOAC (Ed.), 1995. Official Methods of Analysis of AOAC International, Vol. I. Agriculture
Chemicals; Contaminants, Drugs, 16th ed. AOAC International, Arlington, VA.
Cho, C.Y., Slinger, S.J., Bayley, H.S., 1982. Bioenergetics of salmonid fishes: energy in-
take, expenditure and productivity. Comp. Biochem. Physiol. 73B, 25–41.
Commission Directive, 1998. Establishing community methods for the determination of
amino-acids, crude oils and fats, and olanquindox in feeding stuff and amending
Directive 71/393/EEC, annex part A. Determination of amino acids. Offic. J. Eur.
Commun. L257, 14–23.
Hien, T.T.T., Be, T.T., Lee, C.M., Bengtson, D.A., 2015. Development of formulated diets
for snakehead (Channa striata and Channa micropeltes): can phytase and taurine
supplementation increase use of soybean meal to replace fish meal? Aquaculture 448,
334–340.
Hien, T.T.T., Phu, T.M., Tu, T.L.C., Tien, N.V., Duc, P.M., Bengtson, D.A., 2017. Effects of
replacing fish meal with soya protein concentrate on growth, feed efficiency and
digestibility in diets for snakehead, Channa striata. Aquac. Res. 48, 3174–3181.
Hung, L.T., Huy, H.P.V., 2007. Analysis of feeds and fertilizers for sustainable aquaculture
development in Viet Nam. In: Hasan, M.R., Hecht, T., De Silva, S.S., Tacon, A.G.J.
(Eds.), Study and Analysis of Feeds and Fertilizers for Sustainable Aquaculture.
Development. FAO Fisheries Technical Paper. No. 497. FAO, Rome, pp. 331–361.
Iwama, G.K., Tautz, A.F., 1981. A simple growth model for salmonids in hatcheries. Can.
J. Fish. Aquat. Sci. 38, 649–656.
Mercer, L.P., 1982. The quantitative nutrient-response relationship. J. Nutr. 112,
560–566.
Mohanty, S.S., Samantaray, K., 1996. Effect of varying levels of dietary protein on the
growth performance and feed conversion efficiency of snakehead Channa striata fry.
Aquacult. Nutr. 2, 89–94.
National Research Council (NRC), 2011. Nutrient Requirements of Fish and Shrimp.
Animal Nutrition Series. National Academies Press, Washington, DC.
Paripatananont, T., 2002. Snakehead and Pangasius catfish. In: Webster, C.D., Lim, C.E.
(Eds.), Nutrient Requirements and Feeding of Finfish for Aquaculture. CAB
International, pp. 396–401.
Sagada, G., Chen, J.M., Shen, B.Q., Huang, A.X., Sun, L.H., Jiang, J.H., Jin, C.H., 2017.
Optimizing protein and lipid levels in practical diet for juvenile northern snakehead
fish (Channa argus). Anim. Nutr. 3, 156–163.
Samantaray, K., Mohanty, S.S., 1997. Interactions of dietary levels of protein and energy
on fingerling snakehead, Channa striata. Aquaculture 156, 241–249.
Wee, K.L., Tacon, A.G.J., 1982. A preliminary study on the dietary protein requirement of
juvenile snakehead. Bull. Jpn. Soc. Sci. Fish. 48, 1463–1468.
Zehra, S., Khan, M.A., 2012. Dietary protein requirement for fingerling Channa punctatus
(Bloch) based on growth, feed conversion, protein retention and biochemical com-
position. Aquac. Int. 20, 383–395.
Zhang, Y.F., Sun, Z.Z., Wang, A.L., Ye, C.X., Zhu, X., 2017. Effects of dietary protein and
lipid levels on growth, body and plasma biochemical composition and selective gene
expression in liver of hybrid snakehead (Channa maculata ♀ × Channa argus ♂)
fingerlings. Aquaculture 468, 1–9.
Zhao, P.F., Li, F.J., Chen, X.R., Chen, Y.J., Lin, S.M., Zhang, L., Li, Y., 2016. Dietary lipid
concentrations influence growth, liver oxidative stress, and serum metabolites of
juvenile hybrid snakehead (Channa argus x Channa maculata). Aquac. Int. 24,
1353–1364.