Reviews in Fisheries Science & Aquaculture

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A Review on Japanese Eel (Anguilla japonica)

Aquaculture, With Special Emphasis on Nutrition

Ali Hamidoghli, Jinho Bae, Seonghun Won, Seunghan Lee, Dae-Jung Kim &
Sungchul C. Bai

To cite this article: Ali Hamidoghli, Jinho Bae, Seonghun Won, Seunghan Lee, Dae-Jung
Kim & Sungchul C. Bai (2019): A Review on Japanese Eel (Anguilla�japonica) Aquaculture,
With Special Emphasis on Nutrition, Reviews in Fisheries Science & Aquaculture, DOI:
10.1080/23308249.2019.1583165

To link to this article: https://doi.org/10.1080/23308249.2019.1583165

Published online: 19 Mar 2019.

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REVIEWS IN FISHERIES SCIENCE & AQUACULTURE
https://doi.org/10.1080/23308249.2019.1583165

A Review on Japanese Eel (Anguilla japonica) Aquaculture, With Special

Emphasis on Nutrition
Ali Hamidoghlia, Jinho Baea, Seonghun Wona, Seunghan Leea,b, Dae-Jung Kimc, and Sungchul C. Baia
a
Department of Marine Bio-materials and Aquaculture/Feeds & Foods Nutrition Research Center, Pukyong National University, Busan,
Republic of Korea; bUniversity of Idaho, Hagerman Fish Culture Experiment Station, Hagerman, Idaho, USA; cJeju Fisheries Research
Institute, National Institute of Fisheries Science, Jeju, Republic of Korea

ABSTRACT KEYWORDS
Aquaculture has experienced tremendous growth over the last few decades as global pro- Japanese eel; aquaculture;
duction has increased from 16 million tons in 1990 to 110 million tons in 2016. Meanwhile, nutrition; nutritional
the modern aquaculture industry is interested in the production of endangered species with requirements
great economic potential. Japanese eel (Anguilla japonica) is one of the most valuable
species in East Asia with high market demand. But, unfortunately, this fish is critically endan-
gered largely due to excessive harvesting at early life stages and deteriorating environmen-
tal conditions. Therefore, a sustainable aquaculture strategy is necessary to conserve and
rehabilitate Japanese eel stocks while meeting market demand. In order to achieve this
goal, efficient artificial breeding of Japanese eel and knowledge of nutritional requirements
during every life stage is critical. The present review will cover recent research and technol-
ogy in the areas of aquaculture and artificial breeding, and provide comprehensive informa-
tion regarding the nutrition of Japanese eel. The information provided in this study is
intended for researchers working on the aquaculture of Japanese eel by identifying the cur-
rent knowledge and research gaps. This review is also helpful to feed manufacturers in pro-
ducing practical diets for different life stages of Japanese eel.

1. Introduction million tons in 2016. Nowadays, aquaculture produc-

Fish are rich sources of nutrients and contain essential
amino acids and fatty acids (such as long-chain fatty
acids) as well as micronutrients such as vitamins and
minerals. These nutrients, often lacking in many diets
can be found in fish flesh. Therefore, fish consump-
tion aids in the development of organs and prevents
the degeneration of muscle tissue, mitigates the effects
of aging, cardiovascular diseases such as strokes and
myocardial infarctions, neurological diseases, diabetes
and blood pressure (Olsen, 2003; Raatz and Bibus,
2016). This has led to the increase of global fish con-
sumption up to 20 kg per capita as of 2014, which is
16% of all animal proteins (cattle, pork, and chicken)
consumed. Due to the predicted world population
increase to more than 9 billion in 2050, fish is consid-
ered to be one of the main protein sources for satisfy-
ing the forecasted rise demand (FAO, 2016).
Aquaculture has experienced tremendous growth
over the last several decades as the global production
has increased from 16 million tons in 1990 to 110
tion provides about 50% of the global fish supply
while the other half is provided by fisheries (FAO,
2018). Sustainable aquaculture can provide worldwide
nutritional security, reduce the pressure on the envir-
onment and wild fisheries, help with the preservation
and rehabilitation of endangered species, and contrib-
ute to the economic development of countries.
Meanwhile, the modern aquaculture industry is inter-
ested in the production of endangered species with
great economic potential.
Japanese eel (Anguilla japonica) is one of 16 species
of freshwater eel belonging to the Anguoilla genus
(Watanabe, 2003). In East Asian countries such as
China, Japan, and Republic of Korea, A. japonica is
commonly cultured and consumed as a luxury food
item. This fish is also naturally distributed in fresh-
waters of Taiwan, China, Korea, and Northern Japan
(Tesch, 2003). Mitochondrial DNA sequencing of
A. japonica did not show any evidence of genetic
subdivisions between different populations in these

CONTACT Sungchul C. Bai scbai@pknu.ac.kr Department of Marine Bio-materials and Aquaculture/Feeds & Foods Nutrition Research Center,
Pukyong National University, Busan 48513, Republic of Korea.
Color versions of one or more of the figures in the article can be found online at www.tandfonline.com/brfs.
ß 2019 Taylor & Francis Group, LLC
2 A. HAMIDOGHLI ET AL.
countries (Sang et al., 1994; Ishikawa et al., 2001).
Unfortunately, according to the International Union
for the Conservation of Nature and Natural Resources
(IUCN), Japanese eel is listed as endangered under
the red list of threatened species (Jacoby and Gollock,
2014). Currently, the wild population size of this fish
is so low that it is on the brink of extinction.
Although the precise reasons for this decline are
uncertain, some researches have related it to habitat
modifications and impediments to upstream migration
such as dam construction (Itakura et al., 2015).
According to Tatsukawa and Matsumiya (1999) and
Tatsukawa (2003), there is a direct correlation
between the decline in eel catch and population size
to the construction date, height and water volume of
dams. Other reasons such as pollution, disease,
oceanic climate change and habitat losses could also
be involved (Arai, 2016a). But, the reliance of eel
aquaculture on wild-caught juveniles or glass eel
seems to be more significantly influencing the abun-
dance of the Japanese eel population (Tsukamoto,
2013). Also, a great number of Japanese eel popula-
tions are harvested as bycatch in shrimp nets (Lin and
Tzeng, 2010). This has led to the shortage of A. japon-
ica fry and a major problem for farmers of this fish
(Arai, 2014a).
The aforementioned situation highlights the
importance of a standardized aquaculture procedure
for Japanese eel. This can only be achieved by success-
ful artificial breeding technology with minimal
dependence on wild stocks and understanding the
optimal nutritional requirements in order to maximize
production. Therefore, the overall objectives of this
review are to deliver comprehensive information
regarding the nutritional requirements of Japanese eel,
A. japonica while covering the progress in recent
aquacultural and artificial breeding research of this
valuable species.
2. Lifecycle and physiology
In the complex lifecycle of Japanese eel two metamor-
phoses occurs, one in saline waters and the other in
freshwater (Haro, 2003; Rousseau et al., 2012). The
marine stages of Japanese eel are egg and larvae which
are called leptocephalus. The leptocephalus has a
laterally compressed and nearly transparent body,
making it well adapted to marine life (Miller, 2009;
Miller et al., 2015). At this stage, the total length of A.
japonica leptocephali increases from 10.0 to 58.4 mm
with a growth rate of 0.15–0.90 mm day
1 (Arai et al.,
2001). Larval eels undergo metamorphosis and by
developing organs such as intestine, gills and kidneys
they get prepared for migration to freshwaters in early
winter or late spring (Otake, 2003). Kawakami et al.
(2013) were able to link the profiles of thyroid hor-
mone receptors to the developmental stages of
Japanese eel, thereby revealing the extensive role of
thyroid hormones in metamorphosis. Small colorless
eels that arrive in freshwater from the sea are referred
to as “glass eel”, because of their transparent skin. In
the freshwater, glass eel goes through some physio-
logical and morphological changes, such as pigmenta-
tion, and are then called “elvers”. Elvers migrate
upstream with outstanding climbing abilities and turn
into “yellow” eels along the way. With further pro-
gress in the freshwater growing phase, they turn into
“silver” eels which stop eating and growing, then
migrate downstream toward the oceans for reproduc-
tion (Arai et al., 2001; Rousseau et al., 2014). This is
when the second metamorphosis happens, and with
“silvering”, eels become adapted for their transition
to saline waters when sexual maturation begins. The
sexual maturation of Japanese eel will be completed
during the thousands of kilometers they migrate to
reach the waters near the western part of the Mariana
Islands in the Northwestern Pacific Ocean (Tsukamoto
et al., 2003, 2011). It is supposed that adult A. japonica
find their way toward the spawning sites (western part
of Mariana Islands) by reversing the geomagnetic direc-
tion which was followed in the glass eel stage (Nishi
and Kawamura, 2005). Stable isotope analysis by Chow
et al. (2010) showed that Japanese eel will not attempt
feeding during reproductive migration. Chow et al.
(2009) and Tsukamoto et al. (2011) found eggs, larvae
and mature eels near these spawning locations that
helped with the understanding of reproductive charac-
teristics and spawning conditions of this species.
Japanese eel, as is the case with other members of
the Anguilliformes order, have an elongated body, but
have a longer dorsal fin that helps to distinguish it
from other members (Arai, 2016b). The vertebrae
number in this fish is normally 114–118 while this
number for a fish such as common carp (Cyprinus
carpio) is 36–37 (Kottelat and Freyhof, 1972;
Okamura et al., 2002). In contrast to the elongated
body, the Japanese eel has a narrow head that gives it
the ability to burrow inside the sediment of lakes and
rivers and enter small holes in order to search for
food. The operculum is relatively small but the bran-
chiostegal ray, located below the operculum, is well
developed. Therefore, this fish can benefit from
pressure induction during feed intake and suck food
particles or prey inside its mouth which is facilitated

by a set of teeth arranged in the upper jaw
(Alexander, 1970). The gastrointestinal tract of
Japanese eel, as in other physostome fish, has a con-
nection between the esophagus and the swimbladder.
Pyloric ceca do not exist in the gastrointestinal tract
of this fish, such as those that can be found in other
teleost fish. In the larval stage, the stomach can easily
be distinguished from other parts of the alimentary
tract, but when the fish are mature it is hard to differ-
entiate the stomach and gut from one another.
The initiation of first feeding highly depends on
temperature, and at 5  C or 10  C, no feeding is
observed (Fukuda et al., 2009). The digestive enzyme
activity of larvae, on the other hand, is highly depend-
ent on pH, where trypsin, chymotrypsin, amylase and
lipase activity, are higher in pH ranging between 7
and 9 (Murashita et al., 2013). Kurokawa et al. (1995)
described the ontogeny of the Japanese eel larvae
digestive tract during 13 days post hatch (DPH). At 3
DPH the pancreas and liver started to develop, and at
6 DPH the pancreas started to synthesize digestive
enzymes. In the larval stage, the digestive function of
pancreas and digestive enzymes play a major role in
nutrition. After 7 DPH, the anterior mouth opening is
formed, the intestine is developed and larvae are ready
to receive initial feeding. The pancreas probably syn-
thesizes trypsinogen, amylase, and lipase at 8 DPH
with the beginning of exogenous feeding, while digest-
ive capacity progressively increases. It was shown that
after feeding an artificial diet to Japanese eel larvae
the trypsin activity increases by twofold. At the lepto-
cephali stage, the composition of some bacteria and
mucosal cells in the gut and midgut rejected the pre-
vious hypothesis of epidermal nutrient uptake (Hulet
and Robins, 1989; Mochioka and Iwamizu, 1996;
Tsukamoto, 1999). Gut content and composition of
stable nitrogen isotopes of Japanese eel leptocephali
showed evidence of fine detrital particles and aggrega-
tions, with no phytoplankton or zooplankton food
items (Otake and Mochioka, 1994). This shows lepto-
cephali have relatively low nutritional demands, and
they feed on suspended organic matter in the sea,
known as marine snow. During the long metamorphic
period, A. japonica undergoes many changes in body
morphology and digestive tract physiology (Otake,
2003). In the glass eel stage, the mid-gut and stomach
become more developed, along with a change to an
omnivorous regime. Amphipods, copepods, insect lar-
vae such as chironomids, polychaetes, oligochaetes,
and detrital particulate organic matter are considered
as the glass eel diet (Tesch, 2003). Yellow eels are
known to search for any prey that can be ingested in
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 3
the freshwater environment during night time. In this
stage of lifecycle, the olfactory system acts by assisting
the eel in the detection of prey (Moriarty, 2003). In
the silver eel stage, because of the prolonged migra-
tion to saline waters and fasting, the stomach gets
smaller. These morphological changes are in line with
the thickness and thinness of gastrointestinal tract
muscular layer. Generally, the stomach and intestine
in eel are close to one another, and after feed intake,
the stomach becomes expanded and compresses the
intestines. Therefore, continuous feeding seems to be
impossible due to this physiological characteristic. The
eel diet in natural environments includes various
organisms such as small fishes (e.g., Gobiidae), inver-
tebrates (snails and mussels), crustaceans (prawns and
crabs), worms, insect larvae, and zooplankton (Tesch,
2003). The olfactory organ in the paired nostrils plays
an important role in the detection, as well as accept-
ance or rejection of food. At the oropharyngeal and
mouth, there are special taste buds that look like
raised papillae which act as sensory receptors. In bony
fish, taste buds assess the palatability of food and
respond to special prey items or nutritional require-
ments (Caprio et al., 1993; Hunt et al., 2014). Yoshii
et al. (1979), revealed the high electrophysiological
sensitivity of Japanese eel in response to essential
amino acids and carboxylic acids. Further research on
the digestive enzyme activity and changes according
to environmental conditions during all life stages of
Japanese eel is necessary in order to achieve practice
feed formulations.
3. Aquaculture of Japanese eel
Global demand for Japanese eel is mostly met by
aquaculture production with a significant increase in
recent decades. According to FAO (2018), total pro-
duction of Japanese eel in 1991 was 178 thousand
tons that reached 278 thousand tons in 2016
(Figure 1). The price trend of Japanese eel has been
unstable throughout this period due to market
demand and fluctuations in stocks. As it is shown in
Figure 1, an unusual decrease in price happened in
2001 but in recent years the market value seems to be
more stable. In the Republic of Korea, glass eel
imports have experienced a fourfold growth over the
past decade (Table 1), while the export of glass eel has
nearly ceased; likely due to high demands for seed by
the growing eel aquaculture industry in Korea. Also,
significant decreases in silver eel export show growing
domestic consumption (MOF, 2017). Japanese eel is
among the most expensive fish species, making the
 4 A. HAMIDOGHLI ET AL.
300 Production
250
Production (mt × 1000)
200
150
100
50
0 0
1991 1996 2001
Year
Figure 1. Global annual production (metric tons, mt) and
value (dollars) of Japanese eel (Anguilla japonica). Data is
based on FAO (2018).
Table 1. Export and import weight (kg) of glass eel and silver
eel in the Republic of Korea from 2004 to 2016.
Import (kg)
Year Glass eel Silver eel
2004 7,041 5,485,239
2007 5,108 587,043
2010 10,153 3,235,367
2013 54,492 946,933
2016 31,689 988,406
Data is presented from the Ministry of Oceans and Fisheries (MPO 2017).
economic value of eel aquaculture considerable. But
the major impediment to Japanese eel aquaculture is
that artificial reproduction has not been successful at
an industrial level. Nearly all Japanese eel farms in the
Republic of Korea use harvested glass eel from the
wild population for stocking. This is also true for big-
ger producers such as Japan and China. In each coun-
try, dramatic losses in stocks of A. japonica (Figure 2;
FAO, 2018) have resulted in the high price of seed for
farms. On the other hand, the intense competition
between the major A. japonica producers (China,
Japan, Republic of Korea, and Taiwan) has forced
them to lower prices. This has drawn the world indus-
try to the same conclusion; that artificial reproduction
of Japanese eel is the solution.
Generally, any successful aquaculture activity
requires knowledge and control over physiology, arti-
ficial breeding, and nutrition of the target species. The
attempts for artificial reproduction of A. japonica ini-
tiated by Japanese scientists with the successful pro-
duction of fertilized eggs and viable larvae
(Yamamoto and Yamauchi, 1974). These scientists
induced the sexual maturation by injecting salmon
pituitary extract to Japanese eel. Later on, effects of
various types of hormones were tested for in vitro
maturation of oocytes in Japanese eel, where germinal
vesicle break down (GVBD) was considered as an
indicator for maturation. The results of these incuba-
tion techniques showed 17a, 20b-dihydroxy-4-
Value 2500 1400
1200
2000
Value (Million Dollars)
1000
Capture (tons)
1500
800
1000
600
500
400
2006 2011 2016 200
0
1991 1995 2001 2006 2011 2016
Year
Figure 2. Global capture (metric tons, mt) of Japanese eel
(Anguilla japonica). Data is based on FAO (2018).
pregnen-3-one (DHP) was the most potent steroid.
Export (kg) Other steroids such as 17a-OHP, 20b-S and deoxycor-
Glass eel Silver eel ticosterone (4-pregnen-21-o1-3,20-dione) had lower
4,140 38,621 induction potential (Ijiri et al., 1995; Kagawa et al.,
0 62,103 1995; Kim et al., 2016). Ohta et al. (1996), applied
283 460
0 10,785 DHP hormone for maturation and ovulation of A.
138 5,511 japonica and achieved satisfying results by improved
fertility and hatching rates. Later on, larvae of
Japanese eel were reared for two weeks until they
reached 7 mm in total length (Yamauchi et al., 1976).
Survival of normally hatched larvae is highly depend-
ent on rearing conditions, diet and maternal derived
nutritional substances such as protein, fat, carbohy-
drates, vitamins, and hormones in the egg (Adachi
et al., 2003). It usually takes 9 DPH for the Japanese
eel larvae to absorb the yolk sac, and if no external
feeding exists they will grow thin and finally die after
two weeks (Tanaka, 2003). A slurry-type diet made of
powdered shark egg was successfully used for feeding
larvae until they reached 22.8 mm in total length
(Tanaka et al., 2001). The problem was that leptoce-
phali could not grow and water was extremely pol-
luted using a slurry as feed. Increasing physiological
knowledge and investigation of natural spawning
grounds facilitated more commercial success in
Japanese eel aquaculture. Shark egg powder was sup-
plemented with krill, soybean peptide, vitamins, and
minerals which resulted in the cultivation of glass eel
(Tanaka et al., 2003). Artificial breeding of Japanese
eel was accomplished and glass eel was obtained 131
days post hatch (Masuda et al., 2011). Thereafter, Ijiri
et al. (2011) reported the successful production of F2
generation larvae and Kuroki et al. (2014), stated that
the F3 generation is also ready to be produced. But
with low egg quality and extremely low survival rates
of larvae, the production techniques for Japanese glass
eel require more modifications. The progress in artifi-
cial production of Japanese eel was reviewed by

Tanaka (2015) and strong hopes were given to the
development of a stable mass production method for
this species.
The culture method of A. japonica has transformed
from earthen ponds to modern intensive culture and
recirculating aquaculture systems (RAS). But less com-
plex systems with minimal investment are always pre-
ferred. Generally, eels have a great potential for
intensive culture as long as water quality, especially
oxygen, is at an optimal state (Peters et al., 1980). In
pond culture, oxygen can be supplied by any instru-
ment mixing air with water while in RAS, pure oxy-
gen is injected in the culture water. In countries such
as Taiwan, RAS is not popular, and currently surface
waters in most parts of the country are polluted. This
has forced eel aquaculturists to utilize well water;
exceeding the aquafers rate of annual refill (Chen
et al., 2006). The other major problems in eel aquacul-
ture are the physiological deformities occurring at dif-
ferent developmental stages. Deformities such as open
jaw, jaw dislocation, and pericardial edema are occur-
ring at the first feeding and leptocephalus stages.
Several reasons are associated with these deformities
but unfavorable physiochemical conditions such as
water temperature, salinity, and rearing conditions
such as non-standard tanks are the main reasons
(Okamura et al., 2007, 2009). Okamura et al. (2011),
described the notochord abnormalities such as dorsal
curvature in leptocephalus of Japanese eel as one of
the major problems possibly related to an imbalance
in the diet, water chemistry, or temperature. Kim
et al. (2014), demonstrated that water flow and tem-
perature are important in Japanese eel leptocephalus
metamorphosis timing. Nonetheless, eels have an
amazing ability to survive when exposed to air due to
cutaneous respiration (Berg and Steen, 1966). Gousset
(1992), reported that the culture of A. Japonica in a
greenhouse and pond system could result in 8–10 and
1 kg m
2 of production per year, respectively. This is
while water consumption in a greenhouse system is
lower than pond culture. Although, stress in eel farms
is a major concern influencing specific aspects of
physiology, osmoregulation, immune system, repro-
duction, feeding and growth (Wilson, 2014). Stressors
such as environmental factors, starvation or malnutri-
tion, handling or transport and high stocking density
directly impact the brain and result in hormonal
imbalances (Ellis et al., 2012). Kamiya (1972), reported
that ion regulation is directly influenced by cortisol,
increasing branchial Naþ/Kþ-ATPase activity in adult
A. japonica. Okamura et al. (2009), investigated the
effects of salinity on Japanese eel larvae reared in a
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 5
typical hatchery. Results indicated that intermediate
salinity (50% sea water) enhances growth, survival,
and activity of five-day-old larvae. Also, under culture
conditions, the optimal temperature for Japanese eel
was reported to be between 24  C and 28  C and at
temperatures below 12  C feeding was ceased (Usui,
1999). Eggs and larvae of Japanese eel are known to
develop normally at temperatures around 25  C with-
out the occurrence of physiological deformities
(Kurokawa et al., 2008; Okamura et al., 2007; Kim
et al., 2014). Fukuda et al. (2009), examined the influ-
ence of water temperature on otolith growth of A.
japonica, and results showed that under 10  C otolith
growth of glass eel and elvers stopped. All the previ-
ously mentioned studies prove that the efficiency of
aquaculture systems highly depends on the mainten-
ance of water temperature within the optimum rage
of the target species.
4. Feeding and nutritional requirements
In natural environments, the diet of A. japonica both
in freshwater and brackish water stages mostly con-
sists of shrimp, fish, bivalve clams, and crayfish
(Itakura et al., 2015). Due to the special feeding
behavior of Japanese eel, which relies largely on odor,
feed stimulants and attractants play a key role in the
diet. At the glass eel stage, intake of commercial feed
comes with difficulties as physiological regulators of
appetite have not developed properly (Lee et al.,
2003). Attempts to produce practical artificial diets to
support all the nutritional needs of young Japanese eel
have not succeeded. The type of commercial diet used
for juvenile and adult Japanese eel is paste feed that
normally contains 20–45% dry matter. Paste feed is
the mixture of commercial ingredients in powder
form which is mixed with water and oil on farm site.
Extruded and pelleted feeds are also used in some
farms. With the great dependence of Japanese eel sup-
ply on aquaculture, understanding the nutritional
requirements becomes critical. Although, a great deal
of research has focused on the development of opti-
mized artificial diets for cultured Japanese eel, the
industry lacks standard diets covering all economic
and environmental aspects.
4.1. Protein and amino acids
Proteins are the major components of the fish body,
containing more than 65% of its dry weight.
Depending on the sequence of amino acids and the
resulting secondary, tertiary, and quaternary
6 A. HAMIDOGHLI ET AL.

structuring, a vast array of biomolecules and structural Table 2. Essential amino acid requirements (% of dry-matter
components are produced; which is a continual pro- basis) in Japanese eel compared with rainbow trout
(Oncorhynchus mykiss) and channel catfish (Ictalurus punctatus).
cess in living organisms. Due to the necessity of pro-
Japanese eel Rainbow trout Channel catfish
tein, especially in the larval stage of fish, daily Amino acid (%) (Nose 1979) (NRC2011) (NRC2011)
consumption of protein is essential for growth, repro- Arginine 1.7 1.5 1.2
duction, and healing (Webster and Thompson, 2015). Histidine 0.8 0.8 0.6
Isoleucine 1.5 1.1 0.8
The physiology of protein digestion in A. japonica lar- Leucine 2.0 1.5 1.3
vae was studied by Pedersen et al. (2003). In this Lysine 2.0 2.4 1.6
Methionine 1.2 0.7 0.6
study, the augmentation of trypsin activity was clearly Phynylalanine 2.2 0.9 0.7
observed in relation to the increase in age (8–18 Threonine 1.5 1.1 0.7
Tryptophan 0.4 0.3 0.2
DPH) and length of larvae. After day 18, a decrease in Valine 1.5 1.2 0.8
trypsinogen synthesis was observed that could be
related to inadequate artificial feed used in this experi-
Therefore protein-to-energy (P/E) ratio requirement
ment. Pepsin is another critical enzyme for breaking
of juvenile Japanese eel could be 24.1 mg protein kJ
1
down protein molecules. Studies have shown that this
based on growth parameters.
gastric-specific enzyme appears at early stages of glass
It seems that Japanese eel requires all ten essential
eel development, being influenced by the metamor-
amino acids, as the absence of these amino acids in
phosis and changes in dietary habits (Kurokawa et al.,
the diet will cause growth depression. The percentage
2011). In another study by Murashita et al. (2013), it
of amino acid requirements were compared with rain-
was demonstrated that the activity of protein digestive
bow trout (Oncorhynchus mykiss) and channel catfish
enzymes such as trypsin and chymotrypsin is highly
(Ictalurus punctatus) as freshwater carnivorous fish
dependent on water temperature and pH. Therefore,
species and presented in Table 2. Overall essential
it seems that the feed quality and environmental con-
amino acid requirements do not differ significantly
ditions are considerably important in the synthesis
between rainbow trout and Japanese eel, but Japanese
and secretion of protein-digesting enzymes, especially
eel requires more methionine and phenylalanine in
at early life stages, which consequently will influence the diet, whereas differences in essential amino acid
the digestion performance of adults. requirements are more obvious between Japanese eel
As mentioned previously, Japanese eel is a carniv- and channel catfish. Japanese eel requires a higher
orous fish with a high protein requirement. It was portion of each amino acid in the diet comparing to
previously reported that the dietary protein require- channel catfish, especially with regard to isoleucine,
ment of juvenile Japanese eel, as indicated by growth leucine, phenylalanine, threonine, and valine. The
using a purified casein-based diet, is more than 45% main protein sources used in the diet of Japanese eel
(Satoh, 2002). This is comparable to the protein should be able to cover the amino acid requirements
requirement of rainbow trout which is known to be while at the same time have a reasonable price and
42–48% for fingerlings (Hardy, 2002). In a study by good digestibility. Fishmeal has been the main protein
Okorie et al. (2007), dietary protein and protein to source and ingredient in Japanese eel diet (Satoh,
energy ratio requirements of Japanese eel were eval- 2002) because of its well-balanced amino acids, essen-
uated using a 2  3 factorial design. In this study, six tial unsaturated fatty acids, vitamins, and mineral
experimental diets with two protein levels (45 and compositions (Bendiksen et al., 2011). Yet, due to the
50%) and three energy levels (16, 17, and 19 kJ g
1) fast expansion of global aquaculture production,
were used to feed glass eel (0.1 ± 0.02 g) and juvenile increasing demands for this valuable ingredient are
(15.0 ± 3.0 g) Japanese eel. A slurry-type feed was used being frustrated by limited supplies. This has led to
for the glass eel and sinking pellets were used for over-exploitation and high fishmeal prices which
juvenile fish with brown fishmeal and blood meal as eventually brought about the need to replace this
protein sources, fish oil and corn oil as lipid sources, scarce ingredient with more sustainable alternatives
and corn starch and wheat meal to modify the energy (e.g., plant proteins). Probably Lee and Bai (1997),
levels of diets. Results showed no significant differen- started the first attempts in reducing the amount of
ces in weight gain (WG), specific growth rate (SGR), fishmeal in Japanese eel diet. In this study, juvenile
feed efficiency (FE), and protein efficiency ratio (PER) Japanese eel were fed ten isonitrogenous diets with
of Japanese glass eel fed all the experimental diets. decreasing levels of fishmeal being replaced with
Although, juveniles fed diets with 50% protein and blood meal. In higher replacement levels, supplemen-
19 kJ g
1 energy had a higher WG, SGR, and PER. tary amino acids (methionine, isoleucine, and

arginine) were added to the diet to account for the
deficiencies. The results demonstrated that fishmeal
can replace up to 50% and 70% by blood meal with-
out and with the supplementation of essential amino
acids, respectively. Considering the feed conversion
ratio of 1.13 in this study and the amount of fishmeal
(66%) used in commercial Japanese eel diet, fish in-
fish out (FIFO) ratio could be 3.28 for Japanese eel
aquaculture. This is while FIFO ratio for salmon and
shrimp are considered to be 2.27 and 1.36, respect-
ively (Jackson, 2009; Tacona and Metian, 2008). Due
to the aforementioned, future studies should focus on
replacing the amount of fishmeal with sustainable
protein sources.
4.2. Lipid and fatty acids
Classification of lipids can be made either based on
their solubility in organic solvents or their functional
roles. Phospholipids, which are polar lipids, have
structural roles and triglycerides are classified as neu-
tral lipids and responsible for energy storage. Lipids
have critical roles in energy production, cellular struc-
ture, and metabolism regulations. Essential fatty acids
(EFAs) are functional lipids that cannot be synthe-
sized in the body and must be included in the diet in
order to meet the dietary requirements. Therefore,
lipid requirement does not simply mean the percent-
age of fat in the diet, but it refers to all EFAs, sterols,
and phospholipids being provided in a well-balanced
diet. Generally, lipid digestion, absorption, oxidation,
and lipogenesis of fish is similar to mammals. For
example, intestine is the main place of lipid digestion
through the aid of digestive enzymes (lipase and
phospholipase) secreted from the pancreas or hepato-
pancreas. Fatty acids are products of lipid digestion
that are transported by the blood and deposited in the
tissue (Bai, 2012; Tocher and Glencross, 2015).
Carnivorous fish species, such as Japanese eel, can-
not efficiently utilize carbohydrates as an energy
source and thus depend on lipids in the diet to meet
this requirement (Hemre and Deng, 2015). An opti-
mal balance of lipid in the diet can have a protein-
sparing effect with regard to its utilization as an
energy source. In this way, protein can be made avail-
able for other important metabolic activities. For
example, as previously mentioned in the study by
Okorie et al. (2007), 15.3% of mixed oil (5.1% fish oil
and 10.2% corn oil) providing 18.4 kJ g
1 energy
resulted in better growth. Even though fish oil has
been considered as the primary lipid source in
Japanese eel diets, some other studies reported that a
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 7
mixture of cod-liver oil and maize oil (1:2) has
resulted in a favorable growth (Satoh, 2002). More in-
depth lipid and fatty acid requirement studies are
focusing on the EFAs and their deficiency signs on
growth, immunity, and reproduction. Most fish spe-
cies require n
3 and n
6 polyunsaturated fatty
acids (PUFA). In a study by Takeuchi et al. (1980),
young A. japonica were fed different ratios of linoleic
acid (18:2, n
6, LOA) and a-linolenic acid (18:3,
n
3, ALA) which reported that for maximum
growth, diets should include 0.5% of each fatty acid
or 1% of 18:3, n
3. Different oil sources (corn oil
and ollack oil) with variable EFA levels fed to
Japanese eel broodstock demonstrated that PUFA are
necessary for reproduction and growth (Furuita et al.,
2007). The biologically active forms of LOA and ALA
with a higher number of carbons are classified as
highly unsaturated fatty acids (HUFA). These fatty
acids are arachidonic acid (20:4, n
6, ARA), ecosa-
pentaenoic acid (20:5, n
3, EPA) and docosahexae-
noic acid (22:6, n
3, DHA; Tocher, 2010). Bae et al.
(2010), conducted an experiment to evaluate different
levels of dietary ARA in the diet of juvenile Japanese
eel. Six semi-purified diets, using casein, defatted fish-
meal and gelatin with the same levels of protein
(55%) and lipid (15%) were made with 0.07, 0.22,
0.43, 0.57, 0.78, and 1.23% of ARA. After 12 weeks of
feeding trial, diets consisting of 0.57, 0.78, and 1.23%
ARA had the highest WG, SGR, and FE. Broken line
regression analysis also showed that the optimum
dietary ARA requirement of Japanese eel could be
0.69–0.71% of the diet. Japanese eel has the ability for
HUFA biosynthesis from PUFA as proven by the
activity of the key enzymes elongase and desaturase,
which is similar to that of Atlantic salmon, Salmo
salar. But the capacity of biosynthesis depends on
environmental and nutritional factors in the larval
stage (Wang et al., 2014). Shahkar et al. (2016), fed
sub-adult A. Japonica (157 ± 2.5 g) with diets contain-
ing different levels of ARA (0, 0.33, 0.71, 1.06, and
1.65% of diet). Results for WG, SGR, and FE showed
significantly higher values when fish were fed 0.71%
of ARA in the diet. These results were supported by
non-specific immune response analysis such as super-
oxide dismutase and lysozyme activity. The broken
line regression indicated that the optimum ARA
requirement in sub-adult A. japonica could be greater
than 0.71% but less than 0.92% in the diet. Based on
the mentioned studies, ARA was proposed as an
essential fatty acid in Japanese eel diet (Table 3).
There is still much to learn about the quality and
8 A. HAMIDOGHLI ET AL.

quantity requirements of EFA and the biosynthesis
capacities of Japanese eel.
4.3. Vitamins
Vitamins are complex organic compounds needed in
very small quantities that cannot be synthesized
endogenously to sufficient amounts, thus being
required in the diet. They are essential for normal
metabolism, growth, and development. Deficiencies
lead to diseases that are often characteristic of the par-
ticular vitamin in deficit. The general classification of
vitamins is based on their solubility in water or fat.
Fat soluble vitamins are retinol (vitamin A), calciferol
(vitamin D), tocopherol (vitamin E), and menaquin-
one (vitamin K) which are absorbed with dietary fat.
Water soluble vitamins consist of thiamin (vitamin
B1), riboflavin (vitamin B2), niacin (vitamin B3), pan-
tothenic acid (vitamin B5), pyridoxine (vitamin B6),
biotin, folacin, cobalamin (vitamin B12), vitamin C
(ascorbic acid), choline, and myo-inositol (Bai, 2012;
McDowell, 2000). It is well known that environmental
factors (e.g., temperature, salinity, dissolved oxygen,
and pH) and nutrient interactions affect this require-
ment, while the exclusion of any vitamin in fish diet
will produce signs and symptoms of deficiency (Shiau
and Lin, 2015). Furthermore, species type, digestive
physiology, life-stage, and size also have a major influ-
ence on fish vitamin requirements. Simillar to all
other fish species, Japanese eel is susceptible to defi-
ciencies of fat and water-soluble vitamins. Deficiencies
in thiamin, riboflavin, vitamin C, and a-tocopherol
Table 3. Optimum energy (lipid) and essential fatty acid
requirements (% of dry-matter basis) in Japanese eel in differ-
ent life stages.
Optimum level (%) Fish size (g) Reference
Lipida 15.3 15 Okorie et al. (2007)
18:2, n
6 0.5 1.39 Takeuchi et al. (1980)
18:3, n
3 0.5–1 1.39 Takeuchi et al. (1980)
20:4, n
6 0.69–0.71 27 Bae et al. (2010)
20:4, n
6 0.71–0.92 157 Shahkar et al. (2016)
a
Fish oil 5.1% and Corn oil 10.2%
lead to fin and skin hemorrhages as well as other
physical disorders. Vitamins such as pyridoxine,
pantothenic acid, niacin and biotin can also cause
neurological disorders, abnormal swimming and hem-
orrhaging, when not supplemented in sufficient
amounts (Halver, 2002; Satoh, 2002). In addition, vita-
mins play vital roles in the reproductive performance
of adult broodstock Japanese eel. Previously it was pro-
ven by Furuita et al. (2003), that a-tocopherol influen-
ces the egg quality of Japanese eel thus increasing the
fertilization and hatching rate. Later on, a comparison
between vitamin A, C and E content of high and low-
quality eggs in Japanese eel was performed (Furuita
et al., 2009). In this study, high and low quality eggs
were defined as eggs that produce larvae with 80% sur-
vival at 8 DPH and eggs with no hatching, respectively.
Results indicated that high quality Japanese eel eggs
ought to have high levels of vitamin C and E and con-
tain optimal levels of vitamin A.
Quantification of vitamin requirements is essential
to maximize growth and prevent disease, while at the
same time, reduce the costs of practical diets by opti-
mizing vitamin incorporation. Quantitative vitamin
requirement studies for Japanese eel likely began in
the mid-1970s and were continued by other research-
ers (Table 4). Yamakawa et al. (1975), fed Japanese eel
with vitamin-free casein and fatty acid methyl esters
as protein and fat sources, respectively, to understand
the optimum a-tocopherol requirement. Based on
growth, FE and disease signs; the minimum require-
ment of a-tocopherol was concluded to be 200 mg
kg
1. In another study by Bae et al. (2013), juvenile
Japanese eel (15 g) were fed semi-purified diets con-
taining 0, 15, 30, 60, and 120 mg of DL-a-tocopherol.
Five experimental diets were fed two times a day up
to satiation (starting with 3% of body weight and
reducing to 2%) for 12 weeks. Results for growth and
feed efficiency using broken line regression showed
that 21.2–21.6 mg of DL-a-tocopheryl is required per
kg of Japanese eel diet. In a more recent study by
Shahkar et al. (2018), the vitamin E requirement of

Table 4. Quantitative vitamin requirements of Japanese eel at different life stages.

Optimum level
Vitamin (mg kg
1) Fish size (g) Parameters Reference
a-Tocopherol >200 NK Growth, appetite, disease signs Yamakawa et al. (1975)
DL-a-Tocopheryl 21–22 15 Growth, feed efficiency Bae et al. (2013)
a-Tocopherol 213 360 Hematology, tissue concentration, non-specific immune responses Shahkar et al. (2018)
L-Ascorbic acid >27 11 Growth, hematology, serum bacterial activity, tissue concentration Ren et al. (2005)
Ascorbic acida 762 75.5 Liver accumulation, hematology, immune responses Ren et al. (2007)
L-Ascorbyl-2-monophosphate 41–44 15 Growth, feed efficiency Bae et al. (2012a)
L-Ascorbic acid 411–912 360 Tissue accumulation, immune response, gonad histology Shahkar et al. (2015)
NK, not known
a
500 mg kg
1 bovine lactoferrin was also used in the diet

adult Japanese eel (360 g) was investigated based on
hematology, tissue a-tocopherol concentration and
non-specific immune responses. Japanese eel were fed
sinking pellets with 32, 112, 158, 212, and 428 mg of
a-tocopherol per kg of diet. Significant differences in
hemoglobin, white blood cell, plasma glucose, aspar-
tate aminotransferase, alanine aminotransferase, lyso-
zyme, and superoxide dismutase activity along with
broken-line regression for liver a-tocopherol concen-
tration demonstrated that Japanese eel require
212.9 mg kg
1 of vitamin E. Vitamin C requirement
of juvenile Japanese eel was determined in an eight-
week experiment with diets containing 3, 10, 27, 126,
645, and 3135 mg L-ascorbic acid per kg of feed. Based
on growth, liver and brain vitamin C concentrations,
hemoglobin content, serum total protein, and bacterial
activity, more than 27 mg L-ascorbic acid is required
per kg of Japanese eel diet (Ren et al., 2005). Ascorbic
acid in the form of ascorbyl-2-monophosphate and
bovine lactoferrin were incorporated in the experi-
mental diets of juvenile Japanese eel for three weeks.
Evaluation of parameters such as liver ascorbic acid,
hematology and non-specific immune responses
showed that 762 mg ascorbic acid with 500 mg bovine
lactoferrin per kg of feed could be beneficial for
juvenile Japanese eel (Ren et al., 2007). Although, in
this study, growth performance was not evaluated and
only a 3-week feeding trial was conducted. The reeval-
uation of optimum vitamin C requirement in juvenile
Japanese eel (15 g) was performed using 0, 30, 60, 120,
and 1200 mg L-ascorbyl-2-monophosphate in five
semi-purified diets. After a 12-week feeding trial,
results for growth performance, FE, and broken-line
regression demonstrated that the dietary vitamin C
requirement could range from 41.1 to 43.9 mg kg
1 in
the diet of juvenile Japanese eel (Bae et al., 2012a).
Shahkar et al. (2015), evaluated the effects of dietary
vitamin C on tissue concentration, hematology, non-
specific immune response and gonad histology of
male Japanese eel broodstock (360 g). Different levels
of L-ascorbyl-2-monophosphate were added to the
basal diet and were fed (2% of body weight) to
Japanese eel for 16 weeks. The optimal vitamin C
requirement was estimated to be between 410.8 and
911.8 mg kg
1 diet based on broken-line regression
analysis for liver and testes vitamin C concentration.
The aforementioned studies demonstrated how vital
vitamins are for general growth and health of
Japanese eel. The lack of research for other fat and
water-soluble vitamin requirements is a pressing
research need.
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 9
4.4. Minerals
Minerals are inorganic elements required for various
physiological processes in the body. They can be
absorbed into the body through the diet or through
skin or gills from the aquatic environment. Minerals
are classified in two major groups, one of which are
required in relatively large amounts (macrominerals)
and the other group are required in very small
amounts (microminerals). The macrominerals are cal-
cium, phosphorus, potassium, sodium, chloride, and
magnesium while microminerals can include chro-
mium, cobalt, copper, fluoride, iodine, iron, molyb-
denum, manganese, selenium, sulfur, and zinc (NRC,
2011; Webster and Lim, 2015). Research on mineral
requirements come with some difficulties because
some of the minerals are required in such small
amounts that it is hard to make purified diets without
these minerals. It is also difficult to clear the water
body from the investigated minerals, as to prevent
their absorption from the water. It was previously
shown that Japanese eel, as is the case with other fish,
exhibit signs of mineral deficiency. Insufficient cal-
cium, phosphorus, and magnesium in the diet of
Japanese eel leads to reduced growth, poor feed con-
version or anorexia (Lall, 2002). While shortages in
iron could cause hypochromic microcytic anemia in
Japanese eel. It has been mentioned that Japanese eel
require 2700, 400, 2500–3200, and 170 mg of calcium,
magnesium, phosphorus, and iron, respectively, per kg
of diet (Satoh, 2002). Park and Shimizu (1989a),
investigated the quantitative requirements for alumi-
num and iron in Japanese eel. Results for growth and
feed efficiency demonstrate that 15 and 100 mg kg
1
of aluminum and iron, respectively, were the most
optimal amounts for young Japanese eel. They also
reported that 50–100 mg kg
1 of zinc could be benefi-
cial in the diet of Japanese eel (Park and Shimizu,
1989b). There is still a lot to know about the mineral
requirements of Japanese eel in different life stages,
providing many research opportunities in this area.
4.5. Feed additives
Feed additives are ingredients added to aquafeed for
specific purposes other than meeting the nutritional
requirements. The purpose could be an improvement
of the growth and immunity of fish, physical or chem-
ical quality of the feed, quality of the final product
(fish) or the environmental condition. Feed additives
consist of feeding stimulants and palatability
enhancers, antioxidants, pigmentation agents, anti-
microbial agents, organic acids, immunostimulants,
10 A. HAMIDOGHLI ET AL.
Table 5. Administration of different feed additives and their effects on the diet of Japanese eel.
Feed additive Optimum level Fish size (g)
Lactobacillus pentosus 108 CFU g
1 24.63
Bacillus subtilis 108 CFU g
1 8.29
B.subtilisþ MOSa 108 CFU g
1þ5 g kg
1 9.00
Korean mistletoeb 5 g kg
1 200
Propolis 5 g kg
1 7.7
Yellow loess 20 g kg
1 11.5
a
Mannanoligosaccharide.
Viscum album
b
enzymes, and hormones (Bai et al., 2015). Each feed
additive has a specific purpose in the diet, therefore,
they are sometimes referred to as functional
feed additives.
Most of the feed additive research for Japanese eel
has focused on immunostimulants and enhancing dis-
ease resistance in this fish (Table 5). Due to consumer
demands for high-quality fish products and the con-
cern for using antibiotics and drugs in the diet, alter-
native strategies for safeguarding fish immunity are
essential (Kiron, 2012). Probiotics are microbial feed
supplements which can beneficially influence the fish
by producing antimicrobial substances, competing
with harmful bacteria, modulating enzyme activity,
and increasing antibody activity. Their application in
feed can lead to improved growth performance, feed
utilization and immune responses (Iwashita et al.,
2015). The probiotic Lactobacillus pentosus PL11 was
isolated from the intestinal tract of Japanese eel and
supplemented in the diet by Lee et al. (2013). This
study used five experimental diets and challenged
Japanese eel with an etiological agent (Edwardsiella
tarda) that causes several pathogenic symptoms.
Results proved that L. pentosus PL11 improved the
growth and health performance of Japanese eel. In
another study by Lee et al. (2015), sixteen Lactobacilli
species were isolated from the intestine of Japanese eel
and were screened by their ability to produce digestive
enzymes such as amylase, cellulase, protease, and phy-
tase. Among all bacteria, L. pentosus PL11 showed the
highest production of these enzymes, with improved
tolerance against pH and antimicrobial activity against
pathogens. Therefore, this bacterium could be a
potential probiotic in Japanese eel diet. Meanwhile, in
vitro experiments with Japanese eel macrophages, and
the effects of L. pentosus on transcription of cytokine
genes related to immune response (interleukin-1b,
interleukin-6, and tumor necrosis factor-a) showed
that L. pentosus plays an important role in immuno-
modulation (Birhanu et al., 2016). Lee et al. (2017),
isolated Bacillus subtilis WB60 from the intestine of
healthy Japanese eel and compared it with
Lactobacillus plantarum in the diet. Seven
Parameters Reference
Growth, hematology, immune response, challenge test Lee et al. (2013)
Growth, immune response, histology, challenge test Lee et al. (2017)
Growth, immune response, gene expression, histology, challenge test Lee et al. (2018)
Immune response, disease resistance Choi et al. (2008)
Growth, feed utilization, immune response, disease resistance Bae et al. (2012a,b)
Growth, immune response, disease resistance Lee et al. (2017)
experimental diets were prepared by adding three
graded levels of B. subtilis and L. plantarum to the
diet of Japanese eel and comparing them with a con-
trol (no additive) diet. Parameters such as growth per-
formance, feed utilization, non-specific immune
responses and immune related genes were measured.
Also, experimental fish were challenged with Vibrio
anguillarum and anterior intestinal histological exam-
ination was performed. Results indicate that oral
administration of B. subtilis at 108 CFU g
1 could be
an effective probiotic for Japanese eel. Later on, the
same authors incorporated different levels of B. subti-
lis WB60 (0.0, 0.5, and 1.0  107 CFU g
1 diet) and
mannanoligosaccharide (MOS, 0 and 5 g kg
1 diet) in
the diet of juvenile Japanese eel (9 g) and compared it
with dietary oxytetracycline (OTC). Growth perform-
ance, feed efficiency, non-specific immune responses,
immunoglobulin M (IgM) and heat shock protein 70
mRNA expression levels were higher in fish fed diets
containing both the probiotic and prebiotic (ymbi-
otic). Histological analysis, such as intestinal villi
length and muscular layer thickness, showed that
ymbiotic diets resulted in healthier gut for the
Japanese eel. Additionally, probiotic, prebiotic and
ymbiotic diets resulted in higher survival rate when
Japanese eel were challenged with V. anguillarum.
These results showed the synergistic effects of B. subti-
lis WB60 and mannanoligosaccharide in diets of
Japanese eel (Lee et al., 2018). But probiotics are not
the only immunostimulating feed additives success-
fully tested in the diet of Japanese eel. Korean mistle-
toe, a parasitic plant that grows on certain trees, was
shown to have immunostimulating effects and
increased non-specific immune response and disease
resistance of Japanese eel (Choi et al., 2008). Also,
propolis is a compound produced by bees and con-
tains polyphenols, sesquiterpene, quinones, coumarins,
steroids and amino acids (Bonvehi et al., 1994). This
compound was used at 0, 0.25, 0.5, 1.0, 2.0, and 4.0%
of Japanese eel diet by Bae et al. (2012b). Results indi-
cate that 0.5% propolis positively influenced growth,
feed utilization, immune responses and disease resist-
ance of Japanese eel. Yellow loess is a natural mineral

present in mountain soil and contains SiO2 and Al2O3
as the main bioactive compounds (Moon and Jung,
2009). Yellow loess was administered in the diet of
Japanese eel at three levels (5, 10 and 20 g kg
1) and
compared to the most commonly used antibiotic (oxy-
tetracycline). Results showed that 20 g yellow loess per
kg of diet could increase growth, immune response
and disease resistance of Japanese eel and comparable
to diets with oxytetracycline. Therefore, yellow loess
could be a cost-effective feed additive for reducing the
use of antibiotics (Lee et al., 2018). The knowledge of
functional feed additives in Japanese eel nutrition
needs to be further expanded. There are vast catego-
ries of feed additives that have yet to be tested which
may improve the performance and feed quality for
Japanese eel.
5. Conclusions
As was mentioned throughout this review, the devel-
opment of Japanese eel aquaculture highly depends on
performing standard artificial reproduction and recti-
fying the nutritional needs at every life-stage. The
industry should be able to provide farmers with
healthy glass eel and/or juveniles with a reasonable
price without relying on the dwindling wild popula-
tion. Understanding the physiological and environ-
mental needs of Japanese eel reproduction, such as
hormonal changes and physicochemical interaction
with the aquatic environment, could lead to the suc-
cessful establishment of practical hatcheries.
Meanwhile, nutrition plays a vital role in every step of
Japanese eel aquaculture. Providing special feeds to
target all the nutritional requirements of adult and
broodstock fish should lead to the production of
resilient larvae. Increasing survival and optimizing the
growth of Japanese eel larvae can be maximized by
meeting the nutritional demands. At the grow-out
stage, development of practical diets that will reduce
the production costs and at the same time safeguard
growth and immunity is essential. Therefore, future
studies should focus on the replacement of fishmeal
and fish oil by more sustainable protein sources in the
diet, while understanding the nutritional requirements
using recent methodologies such as gene
expression alongside growth and immunity analysis in
Japanese eel.
Funding
This review was supported by the Feed and Food
Nutrition Research Center (FFNRC) at Pukyong
REVIEWS IN FISHERIES SCIENCE & AQUACULTURE 11
National University, Busan, Republic of Korea and
The National Institute of Fisheries Science (NIFS) in
the Republic of Korea (R2018035).
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