Hamidoghlietal 2020-Flounder PDF

Reviews in Fisheries Science & Aquaculture
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Nutrition and Feeding of Olive Flounder

Paralichthys olivaceus: A Review
Ali Hamidoghli, Seonghun Won, Seunghyung Lee, Seunghan Lee, Nathaniel

W. Farris & Sungchul C. Bai
To cite this article: Ali Hamidoghli, Seonghun Won, Seunghyung Lee, Seunghan Lee, Nathaniel
W. Farris & Sungchul C. Bai (2020): Nutrition and Feeding of Olive Flounder Paralichthys�olivaceus:
A Review, Reviews in Fisheries Science & Aquaculture, DOI: 10.1080/23308249.2020.1740166
To link to this article: https://doi.org/10.1080/23308249.2020.1740166
Published online: 18 Mar 2020.
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REVIEWS IN FISHERIES SCIENCE & AQUACULTURE
https://doi.org/10.1080/23308249.2020.1740166
REVIEW
Nutrition and Feeding of Olive Flounder Paralichthys olivaceus: A Review

Ali Hamidoghlia , Seonghun Wona, Seunghyung Leeb, Seunghan Leea, Nathaniel W. Farrisa, and
Sungchul C. Baia
a
Department of Marine Bio-Materials and Aquaculture/Feeds and Foods Nutrition Research Center (FFNRC), Pukyong National
University, Busan, Republic of Korea; bGenetics and Breeding Research Center, National Institute of Fisheries Science, Geojesi,
Republic of Korea
ABSTRACT KEYWORDS
Olive flounder (Paralichthys olivaceus) also referred to as bastard halibut is a marine Aquaculture; olive flounder;
carnivorous species with great commercial value. The production of olive flounder has been feeding; nutrient
dominated by aquaculture with 43 thousand tons of production in 2017 compared to requirements;
Paralichthys olivaceus
fisheries with only 10 thousand tons in the same year. But despite the high market demand
for this species, aquaculture production has not increased accordingly within the last dec-
ade. The development of olive flounder aquaculture has been facing several drawbacks
such as improper feed formulations, dependence on unreliable ingredients (e.g. fish meal
and fish oil), disease outbreaks, and excessive use of harmful antibiotics. All these factors
are major impediments in the sustainable development of olive flounder aquaculture.
A great body of research has been conducted to understand the nutrient requirements
of olive flounder while finding alternative protein sources in the diet. Also, several feed
additives have been successfully tested as immunostimulants in olive flounder. All these
findings are thoroughly discussed in this review, therefore, the information can be utilized
by the industry and academic societies.
Introduction indicates (olivaceus), is usually a dark shade of olive

green, however, as with many flounder species,
The olive flounder (Paralichthys olivaceus), also
known as bastard halibut, Japanese flounder, Nopchi through the modulation of chromatophores in the
(in Korea) or Hirame (in Japan) is a demersal marine skin, pigmentation can vary to a wide degree to pro-
flat fish that is highly valued as food fish in East Asia. vide for camouflage while it lies in wait for prey
It is a species of large-toothed left-eyed flounder (as (Akkaynak et al. 2017). The olive flounder is a carniv-
opposed to the right-eyed Pleuronectidae flounder orous fish and consumes a variety of prey throughout
family) with a demersal/oceanodromous life history. life. The diet of larval and juvenile olive flounder con-
This species is native to the temperate/subtropical sists primarily of mysid shrimp with a shift to larger
coastal waters of the Western Pacific with a preferred prey such as fish as they increase in size. Sexually
depth ranging from 10 m to 100 m (Bai and Lee mature flounder, considered 45 cm in total length,
2010). Olive flounder can handle a wide range of migrate to deeper waters (around 70 m in depth) for
temperature variation between as low as 5.8 C to as spawning from January to May where fertilized eggs
high as 28.5 C, but have an optimal temperature produce larvae that are carried by sea currents to shal-
range of 20–25 C (Iwata et al. 1994). It can reach low sandy bottom nursery grounds. There they
sizes up to 1 m in total length in nature, but are typic- mature for approximately two months before migrat-
ally considered market size between 0.5 kg and to ing offshore to deeper waters in search of prey
1.5 kg (Bai and Lee 2010; Seikai et al. 2010). The cen- (Koshiishi et al. 1991).
ter of distribution is primarily in the temperate waters Wild capture of this species was the primary source
off the southern coasts of Japan and South Korea, but of production during the 80s even though aquaculture
can be found as far south as Northern Borneo and the production in Japan had been under development
Philippines. Olive flounder, as the scientific name since the 1970s. Although aquaculture in Japan and
CONTACT Sungchul C. Bai [email protected] Department of Marine Bio-materials and Aquaculture/Feeds and Foods Nutrition Research Center
(FFNRC), Pukyong National University, Busan 48513, Republic of Korea.
ß 2020 Taylor & Francis Group, LLC
2 A. HAMIDOGHLI ET AL.
wild capture in general have remained fairly stagnant, is not difficult to imagine that biosecurity is very low.
the production technologies that were pioneered in In such situations, antibiotics remain the most sure pre-
1960s Japan would in the 1990s serve as a foundation ventive measure to avoid catastrophic losses. The eco-
for the South Korean aquaculture industry to build nomic impact of disease outbreak in olive flounder
upon. In the year 1991 aquaculture production and culture is not a minor concern. Between 2009 and 2014
capture levels were roughly equal at 8,330 and 8,370 olive flounder production fell nearly 25% from a record
MTs respectively. By the 1992 production in Korea high of 59,328 MT down to 44,740 MT due in large
had surpassed that of Japan, and by the year 2009 part to disease outbreak resulting in as much as a 38%
production in Korea was 54,674 mt and as if 2017 is decline by the year 2017 (Hasan, Jang, Lee, et al. 2018;;
41,207 mt, over 17 times that of Japan which pro- Hasan, Je Jang, et al. 2019) . This highlights the prob-
duced 2,300 MTs that same year (FAO, 2018). lem of biosecurity as aquaculture production rises.
Aquaculture technology, particularly that of closed Many of these problems can be mitigated by advance-
recirculating aquaculture systems (RAS), has seen tre- ments in diet formulations that aim at improving the
mendous advancements in recent years, however, it health status of the fish, such as the myriad of feed
cannot be said this has been the case with olive floun- additives that have been investigated in recent years
der production. Even though production has risen, (Dawood et al. 2018). Many of these additives have
the primary form of production are still land based promising application in bolstering growth and nonspe-
tanks in a flow-through configuration, which are cific immunity, which this review aims at addressing.
located on the coast to exchange water with sea. In Thus an up-to-date review on the nutrition of olive
fact, the water exchange rate in growout facilities can flounder is an important tool in determining the blind-
be as much as 800% daily (Bai and Lee 2010). This of spots in knowledge, and focusing efforts on aspects of
course has obvious impacts on both biosecurity in the olive flounder nutrition that are essential.
facility and the ecology of the receiving waters; hatch-
eries and broodstock facility employ closed system
Nutrient requirements
RAS to address issues with the former.
One of the issues facing the olive flounder aquacul- The nutritional requirements in fish diet are estimated
ture industry is the lack of an affordable and standar- empirically by specifically designed experiments.
dized diet. There is still much to be learned about the Comparisons are made between diets that include dif-
diet of this species. Though aquafeed companies are ferent rates of a specific nutrient with a basal diet that
now starting to tailor commercial feeds to each life contains deficient levels of that nutrient for optimal
stage, farms have traditionally fed there fish a commer- growth, feed efficiency and metabolism (Shearer 2000).
cial pellet for the first couple of months followed by Generally, aquaculture requires a well-nourished and a
raw fish and moist pellets thereafter (Yamashita and high quality feed with reasonable price. All these fac-
Aritaki 2010). Also, most of the trials that have been tors should be considered while formulating diets for
conducted to assess the nutrient requirements in this fish, bearing in mind that different fish species possess
species have been done on larva and juveniles. Of different requirements. Research on nutritional require-
course there are obvious benefits in determining nutri- ments of the olive flounder is still in progress and this
ent requirements at the younger life stages since this is review aims at discussing the available knowledge while
a crucial time in development, but more trials focusing pointing out the research gaps.
on the growout stage will help to guide feed formula-
tors in the development of more and more optimal for-
Protein and amino acids
mulations at all life stages. A major problem in feed
formulation is a high dependence on fish meal (Kader Protein and amino acids play important roles in
et al. 2011; Lee et al. 2012; Niu et al. 2019). Given the almost all critical mechanisms and maintenance of life
increasing scarcity of this finite resource, it is very in all living organisms (NRC (National Research
important that alternative sources of protein be utilized Council) 2011). Fish require protein and amino acids
to improve the sustainability of flounder culture in the for growth, reproduction and survival in different
future as has been the case in other finfish culture. environmental conditions. On the other hand, when
Likewise, antibiotic dependence is a growing concern as dietary protein levels are inadequate in fish feed, it
stocking densities increase. Since most olive flounder causes growth deterioration and poor survival (Wilson
operations are located in densely populated coastal and Halver 1986). Therefore, it is important to iden-
areas consisting of land-based flow through systems, it tify the protein requirement in olive flounder diet,
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 3
Table 1. Dietary protein, amino acids requirements and energy ratio for different stage of olive flounder, Paralichthys olivaceus.
Protein requirement
Fish weight Requirement (%) Energy (KJ g1) Feeding period Ingredients Response criteria1 References
12.5 mg 55.4 18.6 25 day Pollack meal and krill meal RGR Ha et al. 2018
0.3 mg 60 23.3 75 day Flounder muscle WG Bai et al. 2001
4.1 g 46.4-51.2 17 8 weeks Fish meal and casein WG Kim et al. 2002
8.1 g 45 16.7 8 weeks White fishmeal and dextrin WG, FE, SGR and PRE Kim et al. 2004
22.7 g 40-44 19.7 9 weeks White fishmeal and Casein WG, FE, DFI and PER Lee et al. 2002
58.4 g 50.1-59.2 20.7 8 weeks Fish meal and Casein WG Kim et al. 2017
Amino acid requirement
Amino acid Fish weight Requirement Feeding period Response criteria References
Arginine 1.85 g 2.04–2.10% 40 days WG and FCE Alam et al. 2002
(4.10–4.20% of protein)
Lysine 3.0 g 1.5–2.1% 9 weeks WG Forster and Ogata 1998
(3.30–4.60% of protein)
Methionine 2.8 g 1.44–1.49% 40 days WG and FE Alam et al. 2000
(2.88–2.98% of protein)
1
Weight gain ¼ WG, relative growth rate ¼ RGR, feed efficiency ¼ FE, daily feed intake ¼ DFI, protein efficiency ratio ¼ PER, specific growth rate ¼ SGR, pro-
tein retention efficiency ¼ PRE, feed conversion efficiency ¼ FCE.
2
P:E ratio was 27.5 in this experiment.
3
Selectively Bred (F-5).
because not only it affects optimum growth and sur- protein sources, and energy content in feed (NRC
vival, but also the cost of production. Previous (National Research Council) 2011).
research with different life stages of olive flounder The studies for the amino acid requirements in olive
demonstrated that at each stage olive flounder flounder have been carried out for arginine, lysine and
requires a different level of protein in practical diets. methionine (Forster and Ogata 1998; Alam et al. 2000;
Dietary protein and amino acid requirements for olive Alam et al. 2002) and is presented in Table 1.
flounder are shown in Table 1. Bai et al. (2001), Meanwhile, there is a big gap of knowledge in other
observed that optimum requirement of dietary protein essential amino acid requirements in olive flounder.
level in early stage of olive flounder (0.3–330 mg) is Generally it is difficult to perform amino acids require-
around 60% in diet based on growth and survival. In ment studies because there is always some amino acids
another study, Ha et al. (2018), reported that larval in the feed ingredients, thus, purified diets are needed to
olive flounder (12.5–58.6 mg) requires 55.4% protein identify exact amounts of required amino acids in fish
based on relative growth rate (RGR). Kim et al. (2002) (Cowey 1995). Arginine requirements of olive flounder
suggested that diet of juvenile olive flounder (4.1 g) in a purified diet was estimated to be 2.04% and 2.10%
must contain 46.4– 51.2% protein. While growing in diet (or 4.10–4.20 of crude protein) based on percent
weight gain and feed conversion efficiency. Forster and
olive flounder (22.7–110 g) require 40–44% of protein
Ogata (1998), reported that olive flounder required
according to growth and efficient protein utilization
1.5–2.1% (or 3.3–4.6% of crude protein) of lysine in the
(Lee et al. 2002). In a more recent study it was
diets for optimum growth, feed efficiency, and nitrogen
reported that optimum dietary protein in selectively
retention. Furthermore, the optimum dietary level of
bred olive flounder was 50.1–59.2% which is higher
methionine in the presence of 0.06% of dietary cysteine
than normal stocks (Kim et al. 2017). While the
for olive flounder was estimated using break point ana-
selected strain of olive flounder required higher protein
lysis. The values determined based on weight gain and
compared to normal olive flounder, this might be due feed efficiency were 1.49% (or 2.98% of protein) and
to the selective breeding program as it leads to superior 1.44% (or 2.88% of protein) in diet, respectively (Alam
growth rates and is associated with a greater feed et al. 2000). Kim et al. (2017), stated that taurine supple-
intake. Optimum dietary P/E ratio in olive flounder is mentation can improve growth, feed utilization, and
suggested to be 27.5 with diets containing 45% crude innate immunity of olive flounder in low water tempera-
protein and 16.7 kJ g1 energy (Kim et al. 2004). In tures. Although taurine is not an essential amino acid
general, early stages such as larvae or juvenile olive and is synthesized from methionine.
flounder required high protein levels for growth and
survival compared to sub-adult or adult fish (Santos
and Meurer 2018). This is in agreement with previous Lipids and fatty acids
conclusions that dietary protein requirement for opti- Lipids and fatty acid are critical for the functional
mal growth depend on fish size, temperature, quality of integrity of cell membranes and maintaining the
Table 2. Dietary mineral and vitamin requirements of olive flounder, Paralichthys olivaceus.
Mineral requirement
Minerals Fish weight (g) Requirement Ingredient Feeding period Response criteria1 References
Phosphorus 2.0 0.45–0.51% NaH2PO4H2O 8 weeks WG Wang et al. 2005
4.01 0.45–0.57% NaH2PO42H2O 6 weeks WG, FE, SGR, PER, WBP Choi et al. 2005
Ca(H2PO4)H2O
Vitamin requirement
Vitamin Fish weight Requirement Ingredient Feeding period Response criteria References
Vitamin A 1.59 9000 IU/kg Retinyl palmitate 70 days WG Hernandez et al. 2005
Vitamin C 3.0 93 & 150 mg/kg L-ascorbyl-2-polyphosphate 12 weeks WG, TS Wang et al. 2002
Myo-inositol 1.22 800mg/kg Cis-1,2,3,5-trans-4, 26 weeks WG Lee et al. 2006
10 617mg/kg 6-cyclohexanehexol 20 weeks WG Lee et al. 2008
Choline 5.9 847 & 1047mg/kg Choline chloride 8 weeks WG, CC Won et al. 2019
1
Weight gain ¼ WG, feed efficiency ¼ FE, specific growth rate ¼ SGR, protein efficiency ratio ¼ PER, Whole body phosphorus retention, Tissue
saturation ¼ TS, Choline concentration ¼ CC.
structure for normal growth, reproduction and sur- flounder, further studies are needed to determine the
vival of fish (Watanabe 1982). Deficiency of dietary roles of these ingredients in carnivorous fish and
lipid levels in the diet causes the utilization of protein proper amounts in the diet.
for energy purposes which can increase the ammonia
excretion, aggravate water quality and give rise to
Minerals and vitamins
harmful effects on the fish. On the other hand, excess
levels of dietary lipid lead to reduced feed intake and Several studies have been conducted on the role of
growth performance of fish (Peres and Oliva-Teles micronutrients such as minerals and vitamins in olive
1999). In juvenile olive flounder, 46–51% of protein flounder diet. The study of these nutrients is still going
with 6% of lipid in the diet showed optimal growth on and a lot needs to be known on their requirements
and efficient feed utilization (Lee and Kim 2005). and critical roles. According to Cech and Moyle (2000),
According to Jang et al. (2019), juvenile olive flounder vitamins and minerals serve not only for the normal
(3.5–11.6 g) required 1.1–1.25% of eicosapentaenoic cell functions and development but also skeletal struc-
acid (EPA) and 0.75% of docosahexaenoic acid ture, ion regulation and acid/base balance (NRC
(DHA). Additionally, sufficient amounts (16.7%) of (National Research Council) 2011). Dietary mineral
dietary n-3 highly unsaturated fatty acids (n-3 HUFA) and vitamin requirements for olive flounder are pro-
resulted in normal growth and feed efficiency com- vided in Table 2. Wang et al. (2005) reported that opti-
pare to the deficient amounts (1.5%) of n-3 HUFA in mum dietary phosphorus (NaH2PO4H2O) level in
olive flounder. The dietary n-3 HUFA was proven to juvenile olive flounder was between 0.45% and 0.51%
be essential for the optimal growth performance in for maximum weight gain. Additionally, Choi et al.
juvenile olive flounder (Kim, Lee, et al. 2002). (2005) demonstrated that the dietary phosphorus
requirement for olive flounder is 0.45 to 0.57% when it
is supplied as NaH2PO42H2O and Ca(H2PO4)2H2O in
Carbohydrates
the diet. Phosphorus is an inorganic compound of feed
Carbohydrate serve as the part of main energy in the and a key component of energy releasing reactions in
diet for fish as well as increasing the water-stability the TCA cycle. In fish feed, incorporation of bioavail-
and firmness of feed (Lee et al. 2016). Studies regard- able phosphorus sources is as important as optimum
ing requirement or utilization of carbohydrates in amounts (Molina-Poveda 2015).
olive flounder are scarce. Generally, carnivorous fish The optimum dietary vitamin C (L-ascorbyl-2-poly-
such as olive flounder don’t have an efficient mechan- phosphate) ranged from 93 to 150 mg/kg regarding
ism for utilization and digestion of carbohydrate sour- weight gain and tissue saturation. This is while vita-
ces (Krogdahl et al. 2005). The excess dietary min C deficiency in the diet can lead to diseases such
carbohydrates in carnivorous fish can lead to fat as scoliosis and lordosis (Wang et al. 2002). The stud-
deposition by stimulating the activities of lipogenic ies of vitamin C deficiency indicated structural
enzyme (Lin et al. 1978). According to Lee et al. deformities for many fish species such as salmon and
(2003), juvenile olive flounder (4–15 g) showed rainbow trout (Sato et al. 1983; Miyasaki et al. 1992;
improved growth performance and feed utilization Dabrowski et al. 1996). Meanwhile, the deficiency of
when provided diets containing 25% dextrin. Apart vitamin A in fish diet leads to anemia, twisted gill
from the requirement of carbohydrates in olive opercula, poor growth and eye lesions (Poston et al.
Table 3. Animal protein sources as alternative ingredients for fishmeal in olive flounder diet.
Protein source Fish weight Fishmeal level1 Tested levels2 Recommended levels Duration (weeks) References
Meat meal 23 g 60% 0–60% 40% 10 Cho et al. 2005
Meat and bone meal 5g 70% 0–44% 18% 7 Kikuchi et al. 1997
Meat and bone meal 14.5 g 60% 0–20% 20% 6 Lee et al. 2012
Poultry by-product 5.55 g 40% 0–80% 80% 8 Wei et al. 2006
Feather meal 3g 80% 0–80% 20–40% 8 Kikuchi et al. 1994
Tuna by-product 12.6 g 60% 0–100% 30% 7 Kim et al. 2014
Fermented fisheries by-product 5.99 g 50% 0–60% 30% 10 Sun et al. 2007
Silk worm pupae meal 14.5 g 60% 0–20% 10% 6 Lee et al. 2012
1
The level of fish meal in control diet.
2
The levels replacing fish meal in experimental diets.
1977; Villeneuve et al. 2005). Hernandez et al. (2005), protein content, well-balanced amino acid profile,
stated that optimum dietary vitamin A requirement in high digestibility and good palatability (Gatlin et al.
olive flounder could be 9000 IU/kg based on weight 2007). Whereas these properties depend on the source,
gain. Vitamin A is fat soluble-vitamin which plays quality, and species that the FM ingredient was made
crucial roles in epithelial cells, cell differentiation, from. Rahman et al. (2016), compared the apparent
reproduction, mucus production, and vision (NRC digestibility of herring, anchovy, mackerel, sardine,
(National Research Council) 2011). Choline is another tuna, and pollock FM in olive flounder diet.
vitamin that is essential for the component of plas- The results showed significantly higher digestibility
malogens and lecithin in fish. Won et al. (2019), of diets containing mackerel, sardine, and pollock FM.
observed that choline requirements could be between Research on FM replacement in olive flounder feed has
847 and 1047 mg/kg in olive flounder diet based been ongoing for many years and lately has received
on weight gain and liver choline concentrations. more attention as FM replacement is a necessity
Normally, most studies of choline requirements in because of its high market price and limited supply
fish demonstrated a range between 50 and 3000 mg/kg (Hardy 2010). Generally, alternative protein sources are
diet (NRC (National Research Council) 2011). low in one or more limiting amino acids such as lysine
Inositol, sometimes referred to as vitamin B8, is a and methionine compared to FM. Therefore, supple-
carbohydrate that acts as a quasi-vitamin and there- mental crystalline amino acids (AA) have been used as
fore its biologically active form (myo-inositol) is clas- a strategy to meet the AA requirement while reducing
sified as a vitamin in fish nutrition (NRC (National dietary FM (Lee et al. 2019). To date, many studies
Research Council) 2011; Molina-Poveda 2015). Myo- have been performed comparing the efficacy of FM
inositol is a structural constitute of the cell membrane and alternative protein sources and over 30 different
that prevents the accumulation of fat in liver (Cody kinds of protein sources have been proposed as partial
1984). Dietary myo-inositol requirement in juvenile replacements to fishmeal (Tables 3 and 4). One of the
olive flounder (1.2–31 g) was estimated to be approxi- biggest challenges for the aquafeed industry is to fur-
mately 800 mg/kg diet based on growth; whereas ther reduce the amount of FM in feed and increase the
growing olive flounder (10–74 g) required at least number of alternative protein sources with ingredient
617 mg/kg of myo-inositol for growth and lipid diversity in the feed of carnivorous fish (Burr et al.
metabolism (Lee et al. 2006; 2008). These studies 2012) such as olive flounder.
suggested that the requirement of myo-inositol may
change depending on the weight of olive flounder.
Animal protein
Adult olive flounder might be less dependent on
dietary myo-inositol for maintaining normal growth The most commonly used animal protein among FM
and survival (Lee et al. 2008). substitute sources is meat and bone meal, feather meal
and animal/fish by-products. Animal protein has
adequate content of essential amino acids and high
Feed ingredients
digestibility and palatability with minimal anti-nutri-
Across the aquaculture industry, feed costs represent tional factors compared to plant proteins. These ani-
over 50% of operational costs and the protein compo- mal ingredients show tremendous variation in
nent constitutes the highest portion compared to chemical composition which cannot be standardized
other nutrients (Southgate 2003). Among fish feed for feed formulation. These are related to the quality
ingredients included in the olive flounder feed, fish and origin of raw materials and the process involved
meal (FM) is the major protein source due to its high during their production. In addition, animal protein
Table 4. Plant protein sources as alternative ingredients for fishmeal in olive flounder diet.
Protein source1 Fish weight Fishmeal level2 Tested levels3 Recommended levels Duration (weeks) References
SBM 5.0 g 60% 0–20% 20% 8 Choi et al. 2004
SBM 45.5 g 60% 0–30% 20% 10 Choi et al. 2004
SBM 5.0 g 75% 0–55% 45% 8 Kikuchi 1999
SBM 13.2 g 55.8% 0–42% 16% 8 Ye et al. 2011
SBM þ AA 5.0 g 60% 0–40% 30% 8 Choi et al. 2004
SBM þ AA 45.5 g 60% 0–30% 30% 10 Choi et al. 2004
SBM þ AA 3.2 g 60% 0–30% 30% 42 Kim et al. 2008
SBM þ SLP 5.56 g 40% 0–75% 25% 7 Kim et al. 2009
SBM þ CSM þ AA 0.74 g 60% 0–40% 20% 10 Pham et al. 2005
FSBM 8.36 g 70% 0–40% 27.9% 8 Seong et al. 2018
SPC þ AA 2.45 g 74% 0–100% 0% 9 Deng et al. 2006
SBM þ CGM4 6g 75% 0–60% 40% 8 Kikuchi 1999
1
Soybean meal ¼ SBM; amino acid (lysine and Methionine) supplementation ¼ AA; Squid liver powder ¼ SLP; Cottonseed meal ¼ CSM; Fermented soybean
meal ¼ FSBM; Soy protein concentrate ¼ SPC.
2
The level of fishmeal in control diet.
3
The levels replacing fishmeal in experimental diets.
4
Soybean meal ¼ SBM þ Corn gluten meal ¼ CGM þ freeze-dried meat of blue mussel .
sources such as meat and bone meal, poultry by- The anti-nutritional factors in plant sources can cause
product meal or feather meal could have safety lower protein and amino acid utilization, and distal
problems due to the risk of foot-and-mouth disease, enteritis in carnivorous fish such as flounder and
mad cow disease and bird flu that can be transmitted salmonids. For this reason, various physical, chemical,
to humans (Sofos 2008). and biochemical processes including hydrolysis,
Meat meal and meat and bone meal are the most fermentation, and heat-treatments have been devel-
common animal ingredients in flounder feed to replace oped to remove the anti-nutrient factors and enhance
FM. Poultry by-product and feather meal are also utilization of plant ingredients (Storebakken et al.
considered to partially replace FM. These ingredients 2000; Heikkinen et al. 2006).
have proper palatability, high calcium and phosphorus Among the plant protein sources, soybean meal
concentrations, balanced essential AA profile, and (SBM) is the most common ingredient with a more
sufficient amounts of limited AA in plant ingredients favorable protein content and amino acid profile com-
such as lysine, methionine and cysteine. Substitution of pared to other plant meals. Also, SBM is an econom-
FM with animal protein sources, such as meat and ically attractive ingredient that is sustainably produced
bone meal up to 18% and 20% (Kikuchi et al. 1997; (Tangendjaja 2015). The use of fermented plant meal
Lee et al. 2012), meat meal up to 40% without supple- and protein concentrate is increasing as a substitutes
mentation of amino acids (Cho et al. 2005), poultry for FM due to high protein content, low anti-nutrition
by-product up to 80% (Wei et al. 2006) and feather factors, and good digestion. The composition and
meal up to 40% (Kikuchi et al. 1994) has been balance of AAs are inferior to FM and supplemental
successfully done without retardation of growth in olive AAs should be added to the feed in order to maintain
flounder. Also, new alternative sources such as fishery adequate growth as the amount of plant meal and
by-product and worm meal have been recently investi- plant protein concentrate increases (Gaylord and
gated as FM replacers. Tuna by-product and fermented Barrows 2009). Also, it is desirable to use various
fisheries by-product successfully replaced FM up to protein sources together rather than using a single
30% in olive flounder diet (Kim et al. 2014; Yoo et al. source to reduce FM content and balance AAs in the
2007). Lee et al. (2012) reported that dietary silk worm manufacture of commercial feed (Burr et al. 2012).
pupae meal could replace FM without retardation of Several experiments with olive flounder have
growth in olive flounder (Table 3). evaluated SBM as an alternative ingredient for FM
(Table 4). According to these studies, 16–20% and
30% of FM can be replaced with SBM and SBM
Plant protein
with supplemental amino acids, respectively, without
Many different plant-protein sources including reduced growth and feed efficiency ratios (Choi et al.
plant-protein meals and plant-protein concentrates 2004; Kim et al. 2008; Ye et al. 2011). In case of
have been examined in flounder feed (Table 4). processed plant proteins, the substitution of FM with
Hardy (2010), proposed that the global production fermented SBM and corn gluten up to 27.9% and
of alternative ingredients from plant protein sources is 40%, respectively, did not adversely affect growth
sufficient to supply the needs of aquafeeds for future. performance (Kikuchi 1999; Seong et al. 2018). This
is while, soy protein concentrate could not effectively disease. In 2005, along with intensification, water
replace FM protein in diets of olive flounder (Deng et al. quality degradation, and infectious disease outbreaks,
2006). Kim et al. (2009) and Pham et al. (2005) per- South Korean olive flounder farms experienced
formed experiments and formulated diets with a com- approximately 50% mortality in grow-out stage (Kim
bination of SBM and other alternative protein sources to et al. 2009). In addition, one of the reasons for the
replace FM. Substitution of FM with SBM and squid dramatically decreased production of olive flounder
liver powder up to 25% and SBM with cottonseed meal from 2009 to 2017 was believed to be the viral and
up to 20% was possible in olive flounder feed. bacterial infectious diseases (Hasan, Je Jang, et al.
2019). The immediate solution for disease outbreaks
could be the use of antibiotics such as oxytetracycline.
Feed additives
But with global ban on the usage of antibiotic chemi-
There is some debate in the proper definition of feed cals in aquaculture farms, because of the appearance
additives and the ingredients that fall under this classifica- of antibiotic resistance strains, they are no more an
tion. NRC (National Research Council) (2011), defined option for reducing high mortalities in olive flounder
feed additives as nonnutritive components in feed formu- farms (Miranda et al. 2018). An alternative strategy
lations that positively influence the feed or the fish. could be the prevention of disease by improvement of
Whereas, others have defined feed additives as ingredients host resistance and modulation of innate immune
(nutritive/nonnutritive) that are incorporated in the diet responses (Defoirdt et al. 2011). Probiotics are live or
for purposes other than matching up the nutrient require- killed microorganisms incorporated in the diet for
ments such as enhancement of growth, immunity, feed purposes such as improvement of immune system,
quality, and final product properties (Bai et al. 2015; growth performance, and environmental (water qual-
Hamidoghli et al. 2019). All these definitions could be ity) conditions. The mechanism of actions is mainly
correct, considering the fact that aquaculture nutrition is by out-competing pathogenic bacterial, nutrient
an emerging science and every year new functional feed uptake, enzyme production, and enhancement of
additives are introduced to the industry that might not innate immunity (Lee et al. 2017). Prebiotics, on the
match the previous descriptions. Feed additives are a other hand, are non-digestible but fermentable saccha-
diverse group of ingredients used to serve special func- rides providing energy for the selective probiotics. The
tions in the feed formulation. A major group of feed combinations of probiotics and prebiotics in aquafeed
additives are used to improve the performance of target is referred to as synbiotics (Lauzon et al. 2014). The
species by increasing growth, reducing stress, enhancing administration of probiotics, prebiotics, and synbiotics
immune responses, and/or positively affecting water qual- in olive flounder nutrition was extensively reviewed
ity parameters. Other feed additives can be used to by Hasan, Je Jang, et al. (2019) and will be briefly dis-
increase feed quality by stabilizing pellet formation (e.g. cussed in this section (Table 5). Jeong et al. (2006),
binders), increasing preservation period (e.g. antioxidants investigated the effects of Bacillus polyfermenticus,
and antimicrobials), and enhancing the nutrient digestibil- Bacillus licheniformis, and the combination of these
ity (e.g. enzymes). Many of these feed additives have two in the diet (1.0 107 CFU/kg) of olive flounder
brought significant benefits to the aquafeed industry and for eight weeks. The authors concluded that based on
along with the fast growth of global aquafeed production, higher respiratory burst activity and cumulative sur-
the feed additive market is also experiencing an enormous vival percentage, incorporation of these two probiotics
growth (Encarnaç~ao 2016). But the mechanism of action in the diet could be beneficial. Although, in this study,
for feed additives is species-specific and various effects at growth and feed utilization was not significantly influ-
different dosages could be observed based on the physi- enced by the dietary probiotics. On another occasion,
ology and ecosystem of the target species (Valente et al. the dietary inclusion (0.5%) of B. subtilis, B. pumilus
2006). In this section, the application of various functional or B. licheniformis in olive flounder resulted in higher
feed additives will be reviewed on a sea water carnivorous growth, feed utilization, phagocytic activity, and water
species, olive flounder Paralichthys olivaceus. quality indices for those fed by B. subtilis (Cha et al.
2013). B. subtilis is a well-known gram-positive pro-
biotic with vast antimicrobial activities (Todorova and
Probiotics, prebiotics and synbiotics
Kozhuharova 2010) and digestive enzyme induction
Immunostimulants are probably the most investigated capacities (Zokaeifar et al. 2012). Hasan, Jang, et al.
feed additives in olive flounder nutritional studies due (2019), used heat-killed B. subtilis (1 108 CFU/g) in
to high susceptibility of this species to infectious the diet of olive flounder (9.64 g) and observed
Table 5. Dietary Probiotics, prebiotics, and synbiotics as functional feed additives in olive flounder.
Feed additive Amount Fish size (g) Effects Reference
Bacillus polyfermenticusþ 1.0 107 CFU/kg 18 Nonspecific immune responses, cumulative Jeong et al. 2006
Bacillus licheniformis survival percentage
B. subtilis 0.5 % 25 Growth, feed utilization, immune responses, Cha et al. 2013
survival, water quality
Heat-killed B. subtilis 1 108 CFU/g 9.64 Growth, immune responses, disease Hasan et al. 2019
resistance, gene expression
Lactococcus lactis 1 109 CFU/g 80.84 Growth, feed utilization, innate Nguyen et al. 2017
immune responses
Lactobacillus plantarum þ 10 107 CFU/g 37.5 Growth, innate immune response, immune- Beck et al. 2015
L. lactis related gene expression, disease resistance
ß1,3 Glucan 0.1 % 3.2 Growth, feed utilization, innate immune Kim et al. 2006
response, survival,
ß1,3 0.1 % 9.2 Growth, nonspecific immunity Yoo et al. 2007
Glucan þ feed stimulant
GOS1 0.1 % 14 Innate immune responses, disease resistance Hasan, Kim, Jang,
et al. 2018
B. subtilis þ GOS 0.1%þ1 108 CFU/g 10 Growth, Feed utilization, nonspecific Hasan, Jang, Kim,
immunity, histology et al. 2018
FOS þ MOS þB. clausii
1 7
0.25%þ0.25%þ10 CFU/g 21 Growth, digestive enzyme activity, lysozyme Ye et al. 2011
activity, blood biochemistry
1
Gluco oligosaccharide ¼ GOS; Fructo oligosaccharide ¼ FOS; Mannan oligosaccharide ¼ MOS.
significant differences in growth, protein efficiency insignificant differences in growth to the inclusion
ratio, nonspecific immune responses, disease resist- level of glucooligosaccharide, relatively short feeding
ance, and immune-related gene expression (interleu- trial, and the origin of olive flounder seeds. In add-
kin and tumor necrosis factor) between the control ition, these authors stated that the mechanism behind
and probiotic treated groups. Administration of oral administration of glucooligosaccharide and
Lactococcus lactis in the diet of olive flounder resulted increased immune responses are not clear but could
in increased body weight and length, specific growth be related to the promotion of probiotic microorgan-
rate, feed efficiency, and innate immune responses isms in the digestive tract. More clear evidence of the
(Nguyen et al. 2017). Also, growth and immune stim- synergism between glucooligosaccharide and B. subtilis
ulating effects of combined dietary lactic acid bacteria was demonstrated by Hasan, Jang, Kim, et al. (2018).
(Lactobacillus plantarum and Lactococcus lactis) was In this study, oral administration of ß-glucooligosac-
proven in olive flounder by enhanced weight gain, charide (0.1%) plus B. subtilis SJ-10 (1 108 CFU/g)
lysozyme and phagocytic activity, immune related in olive flounder resulted in higher weight gain, spe-
gene expression, and disease resistance (Beck et al. cific growth rate, protein efficiency ratio, respiratory
2015). Selective prebiotics have also shown promising burst, superoxide dismutase activity, serum lysozyme
effects in the diet of olive flounder. ß1,3 Glucan is a activity, myeloperoxidase activity, intestinal microvil-
well-known polysaccharide with vast stimulation of lus length, cumulative survival, and expression of
innate immune reactions such as induction of cellular immune related genes (TNF-a and IL-6). What can be
defense and antimicrobial peptides in vertebrates clearly understood from this study is that supplemen-
(Cerenius et al. 2009). The supplementation of olive tation of the prebiotic or probiotic, individually, did
flounder diet with 0.1% ß1,3 Glucan resulted in not possess the beneficial effects of synbiotic. In a
increased weight gain, specific growth rate, feed effi- complementary study, eight different diets containing
ciency, serum lysozyme activity, and cumulative sur- fructo oligosaccharides (FOS, 0.25%), mannan oligo-
vival (Kim et al. 2006). In a similar study, 0.1% of saccharides (MOS, 0.25%) and Bacillus clausii (1 107
ß1,3 Glucan and a feed stimulant resulted in CFU/g) individually or in combinations were used in
increased growth, feed utilization, hematocrit percent- the diet of olive flounder (Ye et al. 2011). Results
age, chemiluminescent responses, and lysozyme activ- demonstrated that olive flounder fed B.
ity in olive flounder (Yoo et al. 2007). Hasan, Jang, clausii þ FOS þ MOS had significantly improved
Lee, et al. (2018), used 0.1% of ß-glucooligosaccharide weight gain rate, feed conversion ratio, and body pro-
(made from barley b-glucan) in the diet of juvenile tein deposition. This could be partly related to
olive flounder and reported higher super oxidize dis- increased digestive enzyme activities (protease and
mutase and lysozyme activity along with cumulative amylase) of fish fed the synbiotic diet. Also, this diet
survival when challenged with pathogenic resulted in higher serum lysozyme activity along with
Streptococcus iniae. These authors related the depressed triglyceride and cholesterol levels compared
Table 6. Dietary plant extracts, algae, natural minerals, enzyme, and organic acids as functional feed additives in olive flounder.
Feed additive Amount Fish size (g) Effects Reference
Aloe powder 5% 8.1 Disease resistance Kim et al. 2002
Fig and green tea extract 1% 12.9 Growth, feed utilization Cho and Kim 2009
Quercetin 0.5 % 19.9 Growth, immune response, antioxidant capacity Shin et al. 2010
Wood vinegar 0.1 % 3.15 Growth, lysozyme activity, disease resistance Lee et al. 2008
charcoal and wood vinegar mixture 0.5–1 % 14.3 Growth, feed utilization Yoo et al. 2008
Chlorella Powder 2% 1.1 Growth, feed utilization, liver function Koo et al. 2001
Kelp 2–6 % 14.3 Nonspecific immune responses Kim and Lee 2008
Inosine monophosphate 0.1–0.4 % 7.5 Nonspecific immune responses, challenge test Song et al. 2012
Song-Gang Stone 0.5 % 5.0 Disease resistance and anti-mold activities Choi et al. 2004
Seleno-yeast 1.07 mg/kg 4.0 Positive histological responses Bai et al. 2010
Microbial Phytase 0.1–0.2 % 6.15 Apparent digestibility Yoo and Bai 2014
Organic acid blend 4 g/kg 3.5 Intestinal bacterial count, disease resistance Katya et al. 2018
with the control group. From what was mentioned in weight gain, feed efficiency, specific growth rate, anti-
the above studies, probiotics, prebiotics, and synbiotics oxidant capacity, and lysozyme activity in olive floun-
can positively influence the growth and immune der. Wood vinegar is a liquid compound that comes
system of olive flounder (Table 5). Further studies are from the distillation of smoke from burning wood. It
required to identify the natural probiotics in olive contains several phenolic compounds and organic acids
flounder digestive tract and to match up these that can positively influence the digestion and absorp-
probiotics with specific prebiotics for maximum effects. tion of nutrients in animals (Kishi et al. 1999; Li and
Ryu 2001; Choi et al. 2009). Yoo et al. (2008), used
mixture of charcoal and wood vinegar (0.5–1%) in the
Plant-derived feed additives
diet of olive flounder and observed enhanced weight
Plant-derived or phytogenic feed additives are a vast gain and feed efficiency. Quercetin, a plant flavonoid
group of compounds originating form herbs, leaves, found in fruits and vegetables has been used in the diet
roots, fruit, stem, and seeds of different plants (0.5%) of olive flounder (Shin et al. 2010). Growth,
(Table 6). These compounds are incorporated in feed utilization, immunity, antioxidant capacity, and
the diet either as powder form or liquid extracts and physiological homeostasis were improved in olive
can exert growth and immune system enhancements flounder after 60 days of experiment. In another study,
(Encarnaç~ao 2016). The mechanisms of actions different types of plant-source feed additives with high
are related to increased palatability, stimulation of antioxidant activity were used in the diet of olive floun-
digestive enzyme activities, antioxidant and immuno- der (Cho and Kim 2009). After eight weeks of feeding
prophylactic properties, and antimicrobial actions trial, green tea extract (1%) and fig extract (1%)
(Kirubakaran et al. 2010; Chakraborty and Hancz 2011; resulted in higher weight gain and feed efficiency in
Reverter et al. 2014). Aloe vera (Aloe barbadensis) is a olive flounder compared to the control group. These
well-known medicinal herb with large leafs that contain observations could be explained by high antioxidant
a gelatinous compound full of biologically active ingre- and polyphenol levels of green tea and fig (Lim et al.
dients. This feed additive has been previously used in 2005; Paul and Michael 2007).
broiler chicken feed and shows beneficial effects on
intestinal microflora, immune response, and growth
Other functional feed additives
performance (Darabighane and Nahashon 2014). In an
in vitro study with olive flounder kidney phagocytes, Alongside the plant-derived feed additives, macro and
results showed increased respiratory burst activity when microalgae extracts have also been known as effective
1 mg/ml of aloe was incubated for 24 hr (Kim et al. feed additives in aquaculture due to rich sources
1999). This could be an evidence for the immunosti- of amino acids, polysaccharides, vitamins and poly-
mulating effects of aloe as feed additive. Later on, Kim phenols (Dawood et al. 2018). Application of algae
et al. (2002), performed a feeding trial and used 5% of feed additives in fish diet have shown positive effects
aloe powder in the diet of olive flounder. These authors on growth, feed utilization, lipid metabolism, stress
reported higher chemiluminescent response and disease and disease resistance, immune responses, and liver
resistance against Edwardsiella tarda bacteria in aloe- function (Bai et al. 2001; Dawood et al. 2018). The
fed fish. Also, dietary wood vinegar (0.1%) was shown widely distributed Chlorella is one of well-known
to increase the cumulative survival of olive flounder microalgae that can exist in autotrophic, heterotrophic
challenged with E. tarda (Lee et al. 2008). These and mixotrophic conditions, and therefore easily
authors reported that this feed additive enhanced cultured (Ahmad et al. 2018). Dried powder Chlorella
was used in olive flounder diet (2%) and was reported and forms selenomethionine which is stored in protein
to significantly improve weight gain, feed efficiency, as an organic form of selenium (Bierla et al. 2012).
and liver condition (Koo et al. 2001). On the other Organic selenium is fundamental for the functioning of
hand, dried powder kelp macroalgae (Ecklonia cava) selenoproteins that play important roles is antioxidant
at 2, 4, or 6% did not positively influence growth per- defense, immune responses, and cell signaling (Pacitti
formance of olive flounder but enhanced nonspecific et al. 2015). Katya et al. (2018), fed olive flounder with
immune responses such as activated neutrophils, lyso- organic acid blends (4 g/kg formic acid, ammonium
zyme activity, and myeloperoxidase activity (Kim and formate, and propionic acid). Results did not show sig-
Lee 2008). nificant changes in growth but demonstrated significant
Several other functional feed additives, with different differences in cumulative survival with the control
origins, have been successfully used in olive flounder group. In another study by Yoo and Bai (2014), dietary
diet. Song et al. (2012), used 0.1–0.4% of inosine mono- microbial phytase (0.1–0.2%) was shown to improve
phosphate in the diet of olive flounder and observed apparent digestibility of olive flounder fed a low fish
increased nonspecific immune responses (myeloperoxi- meal diet. The mechanism of action is by increasing
dase and lysozyme activity) and disease resistance availability of phosphorus content in olive flounder
against Streptococcus iniae. Inosine monophosphate is a feed by converting indigestible phytate phosphorus to
nucleotide with various key roles in biochemical proc- bioavailable phosphorus.
esses such as cell signaling pathways and serving
nutrients for biosynthesis (Asaduzzaman et al. 2017).
Feeding strategy
Song-gang stone is a natural mineral that mainly con-
tains SiO2, Al2O3, Fe2O3, CaO, K2O and Na2O (Won Development of feeding strategies optimal for culture
et al. 2017). Olive flounder fed 0.5% of dietary song- of a desired fish species is critical for the success of
gang stone showed significantly increased disease resist- aquaculture operation because feeds generally account
ance against Edwardsiella tarda while this feed additive for more than half of total production costs
was proven to have anti-mold activities (Choi et al. (Southgate 2003). Manufacturing a nutritionally bal-
2004). Selenium-enriched yeast (seleno-yeast) has also anced and cost-effective diet should precede feeding
been tested in the diet (1.07 mg/kg) of olive flounder by strategies, which is discussed throughout this review
Bai et al. (2010) and resulted in positive histological paper. Feeding rate (i.e., feed ration), a major compo-
responses. Cultured yeast in selenium-rich medium nent of the feeding strategies being investigated in the
absorbs selenium that replaces the sulfur in methionine flounder is reviewed in this section.
Figure 1. Theoretical relationship between specific growth rate, feed conversion ratio, and feeding rate in fish (modified from De
Silva and Anderson 1995). An optimum feeding rate can be defined as (1) the rate at which efficiency of feed utilization is max-
imum (A) or (2) the rate at which growth is maximum (B).
Table 7. The summary of feeding rate studies conducted on olive flounder (modified from Lee et al. 2019).
Fish Size1 (g) Temperature ( C) Feeding rate (% BW/d) Estimated OFR2 (% BW/d) Reference
0.27–0.69 16.6–18.3 1–16 5.9–13.5 Lee, Lee, et al. 2018
2.13–4.71 17.1 ± 0.4 1–11 5.7–10.2 Lee et al. 2019
3.6–14 22.3–24.7 1–2.8 2.6 Kim et al. 2007
5–9.1 20 ± 1 0–5.52 5.17 Okorie et al. 2013
5.5–15.7 25.7 ± 0.5 0–5.02 4.82–6.36 Kim, Hwang, et al. 2011
7.7–11 14.8 ± 0.3 0–2.84 1.97–2.51 Kim, Hwang, et al. 2011
14.8–21.2 24.1 ± 3.4 1–6 4.6–8 Lee et al. 2019
15.7–20.2 16–17 0–2.7 2.42 Kim, Kim, et al. 2011
20.2–28.9 20 ± 1 0–4.12 3.47 Okorie et al. 2013
240.4–288.3 19–21 0–1.25 1.09 Lee et al. 2014
316.7–372.5 21–24 0–1 0.99 Oh et al. 2014
370–412.8 12–14 0–0.75 0.51 Kim Kim, Kim, Lee,
Han, Lee 2014
384.2–431.6 20–24.5 0–0.9 0.74 Kim, Kim, Kim, Lee,
Han, Kim, et al. 2014
1
Range between mean initial and final body weights when the feeding trial was conducted.
2
Optimum feeding rates (OFR) estimated by either the ANOVA or regression model analysis, including one-slope straight broken-line, two-slope straight
broken-line, quadratic broken-line or quadratic models.
An optimum feeding rate can be defined as 1) the is agreeable to findings reported within the same species
rate at which efficiency of feed utilization is maximum (Lee, Lee, et al. 2018) and in other fish species such as
or 2) the rate at which growth is maximum sturgeons (Lee et al. 2016; Lee, Zhao, et al. 2018).
(Figure 1). It is obvious that the former has an eco- Feeding fish at its level that maximizes gain in
nomic advantage in saving feed costs, while the latter weight (i.e., ad-libitum feeding) has been often prac-
can shorten the grow-out period so that reduce other ticed by researchers and farmers. Sometimes, ad-libi-
costs associated with culture (De Silva and Anderson tum feeding may cause overfeeding because one cannot
1995). The latter has been more often accepted by fish surely decide when to stop feeding. A recent study con-
nutritionists (e.g., Deng et al. 2003; Lee et al. 2014). A ducted on the flounder evaluated how various feeding
brief summary of the feeding rate studies conducted rates influenced immunity (Lee, Lee, et al. 2018).
on the flounder is provided in Table 7. Various fish Interestingly, the expression of immune-related genes
sizes ranging from 0.27 g to 432 g and a wide range of in whole body was significantly downregulated in the
water temperature from 14.8 to 25.7 C have been flounder fed at the feeding rate of 10%, 13%, and 16%
tested in the previous studies. The optimum feeding BW/d in comparison to that shown in the flounder fed
rates for the given experimental conditions (e.g., fish at the feeding rate of 7% BW/d. According to the opti-
mum feeding rate of 11.9% BW/d for the flounder esti-
size, water temperature, feed composition, etc.) were
mated by the best-fit model, the feeding rate of 13%
estimated by either analysis of variance (ANOVA) or
and 16% BW/d was considered as overfeeding.
regression model analysis. The feeding table for feed-
Although the feeding rate of 10% BW/d was below the
ing the flounder in various sizes (5–600 g) and tem-
estimated optimum feeding rate, this sub-optimum
peratures (15–25 C) is provided in Table 8.
feeding rate still resulted in alteration of the immune-
It is worth to point out that selection of an appro-
related gene expression. Taken together, feeding a cul-
priate statistical model that best fits datasets obtained
tured fish seems to be a sophisticated job and careful
from conducting a growth trial is critical. A recent
management in feeding strategy should be considered
study conducted on the flounder demonstrated that in order for the success of aquaculture operation.
the test of various regression models, including one-
slope straight broken-line, two-slope straight broken-
line, quadratic broken-line, and quadratic models, Conclusions
resulted in variable estimates (Lee et al. 2019). By the With the global expansion of aquaculture, market
analysis of model selection criteria (e.g., coefficient of demands for commercial fish species such as olive
correlation and corrected Akaike information criter- flounder will continue to increase. Olive flounder
ion), the quadratic broken-line model was turned out aquaculture is a growing industry that relies, to a high
to be the best-fit model among the tested models (Lee extent, on proper feed at different life stages.
et al. 2019). This model describes that the rate of Understanding all the essential nutrient requirements
growth decreases as the feeding rate increases up to such as amino acids, fatty acids, vitamins and miner-
the optimal level, then the growth response plateaus als are a prerequisite for successful olive flounder
beyond its level. The outcome shown in Lee et al. (2019) mass production. In addition, according to the future
Table 8. Estimated optimum feeding rates (% body weight per day) at below or optimum water tempera-
tures for olive flounder1.
Below optimum temperature ( C) Optimum temperature ( C)
Body weight (g) 15 17 19 21 23 25
10 1.72 2.78 3.22 3.31 3.68 3.46
15 1.50 2.39 2.80 2.97 3.33 3.11
20 1.36 2.14 2.53 2.75 3.10 2.87
25 1.26 1.97 2.34 2.59 2.93 2.71
30 1.18 1.84 2.20 2.47 2.80 2.58
35 1.12 1.73 2.08 2.37 2.69 2.47
40 1.07 1.65 1.99 2.28 2.61 2.39
45 1.03 1.57 1.91 2.21 2.53 2.31
50 0.99 1.51 1.84 2.15 2.46 2.25
60 0.93 1.41 1.73 2.05 2.35 2.14
70 0.89 1.33 1.64 1.96 2.27 2.05
80 0.85 1.27 1.56 1.89 2.19 1.98
90 0.81 1.21 1.50 1.84 2.13 1.92
100 0.79 1.16 1.45 1.78 2.07 1.86
150 0.68 1.00 1.26 1.60 1.87 1.67
200 0.62 0.90 1.14 1.48 1.74 1.55
250 0.58 0.82 1.05 1.39 1.65 1.46
300 0.54 0.77 0.99 1.33 1.57 1.39
350 0.51 0.73 0.94 1.27 1.51 1.33
400 0.49 0.69 0.89 1.23 1.46 1.28
450 0.47 0.66 0.86 1.19 1.42 1.24
500 0.46 0.63 0.83 1.16 1.39 1.21
550 0.44 0.61 0.80 1.13 1.35 1.18
600 0.43 0.59 0.78 1.10 1.32 1.15
1
The values were adopted from the website (nifs.go.kr/fishfeed/view/supply/Calc.jsp) available for public use.
direction of modern aquaculture, investigating the biochemical parameters. Aquaculture. 205(1–2):127–140.

nutritional requirements of this species in controlled doi:10.1016/S0044-8486(01)00670-6
Akkaynak D, Siemann LA, Barbosa A, M€athger LM. 2017.
systems such as RAS is a priority. In these systems, a
Changeable camouflage: how well can flounder resemble
high level of attention to detail in factors other than
the colour and spatial scale of substrates in their natural
growth such as water quality parameters is required. habitats? R Soc Open Sci. 4(3):160824. doi:10.1098/rsos.
Sustainable development of olive flounder aquaculture 160824
relies on standardized diets that are economically feas- Asaduzzaman M, Ikeda D, Abol-Munafi AB, Bulbul M, Ali
ible and environmentally responsible. ME, Kinoshita S, Watabe S, Kader MA. 2017. Dietary
supplementation of inosine monophosphate promotes
cellular growth of muscle and upregulates growth-related
Funding gene expression in Nile tilapia Oreochromis niloticus.
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This review was supported by the Feeds and Foods
10.033
Nutrition Research Center (FFNRC), Pukyong
Bai SC, Cha YT, Wang X. 2001. A preliminary study on the
National University.
dietary protein requirement of larval Japanese flounder
Paralichthys olivaceus. N Am J Aquacult. 63(2):92–98.
ORCID doi:10.1577/1548-8454(2001)063<0092:APSOTD>2.0.
CO;2
Ali Hamidoghli http://orcid.org/0000-0003-1745-3690
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Reviews in Fisheries Science & Aquaculture

ISSN: 2330-8249 (Print) 2330-8257 (Online) Journal homepage: https://www.tandfonline.com/loi/brfs21

Nutrition and Feeding of Olive Flounder

Ali Hamidoghli, Seonghun Won, Seunghyung Lee, Seunghan Lee, Nathaniel

To link to this article: https://doi.org/10.1080/23308249.2020.1740166

Published online: 18 Mar 2020.

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Nutrition and Feeding of Olive Flounder Paralichthys olivaceus: A Review

Introduction indicates (olivaceus), is usually a dark shade of olive

direction of modern aquaculture, investigating the biochemical parameters. Aquaculture. 205(1–2):127–140.

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