Abdel-Rahim Et Al-2019-Journal of The World Aquaculture Society

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To what extent can maternal inherited immunity acquired from a crustacean-

enhanced diet improve the performance and vitality of the offspring and
enhance proﬁtability of European...
Article in Journal of World Aquaculture Society · February 2019

DOI: 10.1111/jwas.12598
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Received: 19 May 2018 Revised: 11 January 2019 Accepted: 27 January 2019
DOI: 10.1111/jwas.12598
APPLIED STUDIES
To what extent can maternal inherited immunity

acquired from a crustacean-enhanced diet
improve the performance and vitality of the
offspring and enhance profitability of European
Sea bass (Dicentrarchus labrax)?
Mohamed M. Abdel-Rahim1 | Abdallah T. Mansour2 |
Mohamed H. Mona3 | Mona M. El-Gamal3 | Mohamed M. El Atafy4
1
Division of Aquaculture, National Institute of
Oceanography and Fisheries (NIOF), This study was conducted to investigate the effects of maternal
Alexandria, Egypt inherited immunity acquired from crustacean-enhanced diets on
2
Animal and Fish Production Department, the vitality and profitability of sea bass offspring. Newly hatched
Faculty of Agriculture (Saba Basha), Alexandria
larvae produced from three groups of broodstock were evaluated.
University, Alexandria, Egypt
3
The broodstock were fed (a) a basal diet (BD), (b) a Palaemon-
Zoology Department, Faculty of Science,
Tanta University, Tanta, Egypt supplemented diet (PSD), and (c) an Artemia-supplemented diet
4
General Authority for Fish Resources (ASD) for 42 days. A total of 400,000 larvae at 3 days posthatch
Development (GAFRD), Alexandria, Egypt (DPH) produced from each treatment were stocked in larval rearing
Correspondence tanks at 40 larvae/L for 42 days. Survival (%) was improved by
Mohamed M. Abdel-Rahim, Division of 37 and 9.96% in the groups fed ASD and PSD compared with the
Aquaculture, National Institute of
control group. The growth, swim bladder (%), and condition factor
Oceanography and Fisheries (NIOF),
Alexandria, Kayet Bey, Anfoushy P.O., all significantly (p ≤ 0.05) improved in the postlarvae produced from
Alexandria, Egypt. broodstock enhanced with crustacean diets. Compared with the
Email: mohamed_m_ar@yahoo.com
BD group, the serum lysozyme activities of the fish groups fed ASD
and PSD increased by 45.6 and 11.7%, respectively. Sea bass fry
(90DPH) produced from broodstock fed ASD showed the best tol-
erance to salinity/temperature stress tests. Furthermore, the profit-
ability improved in ASD and PSD compared with the BD group. In
conclusion, sea bass broodstock enhanced with Artemia biomass
produced offspring of superior quality with less cost and greater
profit margins.
KEYWORDS
artemia, Dicentrarchus labrax, fry performance, lysozyme activity,

palaemon, profitability, sea bass, stress tolerance
© Copyright by the World Aquaculture Society 2019

J World Aquacult Soc. 2019;1–25. wileyonlinelibrary.com/journal/jwas 1
2 ABDEL-RAHIM ET AL.
1 | I N T R O D U CT I O N
The European sea bass (Dicentrarchus labrax) is one of the most important commercial fish widely cultured in the
Mediterranean region. It ranks second after Atlantic salmon not only with regard to commercial cultivation but also
as the most popular seafood in Europe (FAO, 2012). Compared with the global aquaculture production of sea bream,
which reached 185,980 tons in 2016, the production of sea bass reached 191,003 tons (FAO, 2018), and this produc-
tion is still rapidly expanding throughout the Mediterranean countries (Basurco & Abellán, 1999). Egypt is one of the
main sea bass producers, with an annual production of 24,498 tons in 2016 originating from fish farms (FAO, 2018).
However, the exponential growth of sea bass farming in Egypt still faces several obstacles. One of the main chal-
lenges is the low quality of hatchery-produced fry compared with wild sea bass; this discrepancy is attributed primar-
ily to the low quality of broodstock nutrition (Essa, 2013; Srour, Essa, Abdel-Rahim, & Mansour, 2016).
Undoubtedly, broodstock nutrition is one of the most vital research areas of fish nutrition, yet it still requires a
more thorough understanding. An improvement in broodstock nutrition has been shown to greatly improve not only
the egg and sperm quality but also the seed production (both the quantity and the quality). Numerous studies have
clarified and confirmed an improved fry quality through the enhancement of broodstock nutrition (Abdollahi, Hei-
dari, & Aghamaali, 2016; Abrehouch, Ait, Chebbaki, Akharbach, & Idaomar, 2010; Aby-ayad, Melard, & Kestemont,
1997; Dhert et al., 2004; Ershad, Mousavi-Sabet, Falahatkar, & Moradkhani, 2009; Fernández-Palacios, Norberg,
Izquierdo, & Hamre, 2011; Furuita, Tanaka, Yamamoto, Suzuki, & Takeuchi, 2003; Izquierdo, Femandez-Palacios, &
Tacon, 2001; Wang & Zhang, 2010; Zhang, Wang, & Wang, 2013).
It is evident that many of the problems encountered by newly hatched larvae during the early rearing stages are
directly related to the broodstock feeding regime (i.e., the feed quality, quantity, and duration). For instance, an
increase in n-3 highly unsaturated fatty acids (HUFA) in broodstock diets, particularly their docosahexaenoic acid
(DHA) levels, was reported to significantly enhance the weight of fish larvae and their resistance to osmotic shock
(Aby-ayad et al., 1997).
The most vital issue regarding the correlation between the broodstock nutrition and larval quality is maternal
immunity. In the early stages of fish larvae, the high mortality rate attributed to infectious diseases resulting from a
limited self-defense mechanism (Mulero, García-Ayala, Meseguer, & Mulero, 2007) causes financial losses in the
Western European aquaculture sector to reach 10% (Dalmo, 2005). The early stages of fish life rely mainly on the
immunological substances transferred from broodstock to offspring, which is referred to as maternal immunity
(Meijer et al., 2004; Mulero et al., 2007). Therefore, mass mortality in the early phases of fish life could be prevented
via the vertical transmission of an innate resistance against infection (Swain and Nayak, 2009).
The broodstock feeding regime usually consists of two major portions, namely, dry diets and natural, fresh food.
The replacement of fresh food with dry feed (DF) is a progressive global trend (Wouters, Lavens, Nieto, & Sorgeloos,
2001) because the use of dry diets over fresh food possesses several advantages, for example, dry food can ensure a
continuous supply; it has a high quality; it is easy to produce and use as well as store; and it is suitable for the delivery
of chemotherapeutics, immune stimulants, and hormones; moreover, it can help maintain water quality and reduce
the spread of infectious diseases (Harrison, 1990, 1997). There have been many attempts to improve the quality of
dry broodstock feed. One such example is the incorporation of vitamin C supplementation, which increases the lyso-
zyme contents of fertilized eggs, embryos, and larvae of D. labrax (Cecchini, Terova, Caricato, & Saroglia, 2000).
According to several researchers, crustaceans such as Artemia and shrimp biomass can provide extra benefits;
accordingly, they have been used for the induction and reinforcement of sexual maturation and to increase fertiliza-
tion rates (Naessens et al., 1997; Wouters et al., 1999). The addition of frozen Artemia biomass to dry broodstock
feed can increase diet ingestion and stimulate ovarian maturation (Wouters et al., 2000). Moreover, Naessens et al.
(1997) and Wouters et al. (1999) reported that female maturation, spawning frequency, and larval quality were
improved through the use of Artemia biomass mixed with DF. The nutritional value of on-grown and adult Artemia is
superior to that of newly hatched nauplii as they have a higher protein content and are richer in essential amino acids
ABDEL-RAHIM ET AL. 3
(Dhont & Sorgeloos, 2002; Lim, Soh, Dhert, & Sorgeloos, 2001; Sorgeloos, Dhert, & Candreva, 2001). In addition,
Artemia biomass can be easily collected from natural lakes or artificially produced in tanks (Dhont & Sorge-
loos, 2002).
Decapod shrimps, Palaemon spp., are a coastal species of shrimp found widely throughout the world, including
the Mediterranean, Black Sea, and Atlantic Ocean (Kelly, Tully, Lehane, & Breathnach, 2009). In addition, Reis et al.
(2015) reported that feeding on crustacean decapods (Palaemon elegans and Grapsus adscensionis) improved the
growth and survival of octopus paralarvae. However, no available literature has presented the effects of feeding
Palaemon to marine broodstock. Therefore, a comprehensive study consisting of two parts was carried out to evalu-
ate the effects of crustacean-enhanced maturation diets on the overall performance of a sea bass hatchery. The first
part focused on the reproductive performance, while the second part focused on the vitality of the offspring. The
current study (Part 2) aimed to evaluate the effects of crustacean-enhanced dry broodstock feeds containing both
Artemia biomass and Palaemon biomass on the efficiency of sea bass (D. labrax) during their larval/postlarval rearing
stages in terms of their survival, growth performance, swim bladder inflation, salinity/temperature stress resistance,
maternal inherited immunity, and economic performance. The price of fingerlings has been a critical variable cost in
the sea bass mariculture industry; therefore, any reduction in the price of the offspring that can be grown out suc-
cessfully to a marketable size is required and deserves more attention from scientific research. Accordingly, the
objective of this economic assessment is to answer the following questions: (a) are the tested feed additives (Artemia
and Palaemon-biomass) likely to have a positive role in the profitability? and (b) To what extent can the quality of
maturation diets affect the economic viability of marine hatcheries?
2 | MATERIALS AND METHODS
2.1 | Broodstock conditions

Figure 1 illustrates the graphical design of the comprehensive study (Part 1 and 2). The sea bass (D. labrax) brood-
stock were 3 years old, farm-reared, and produced from parents captured from the Mediterranean Sea off the coast
of the Damietta Governorate, Egypt. The mean fish weight was 1.36 ± 0.06 kg/female and 1.01 ± 0.11 kg/male. The
fish were distributed into three identical 50-ton cement broodstock maturation tanks representing the three tested
FIGURE 1 Graphical design of the experiment

diets at a ratio of three females to two males (with a total of 10 fishes in each tank). The experiment was carried out
in the Marine Finfish Hatchery, Kilo 21, General Authority for Fish Resources Development (GAFRD), Alexandria,
Egypt. Each tank was supplied with well-aerated running seawater from the Mediterranean Sea, passed through a
sand filter and exposed to a natural photoperiod. At this latitude, the photoperiod reached a maximum of 11 hr of
light and 13 hr of dark from the 10th of January to the 21st of February 2016. The fish were maintained under a nat-
ural temperature regime throughout the experiment (15 ± 2 C); the pH ranged between 7.8 and 8.0, the salinity ran-
ged between 34 and 36 ppt, and the total ammonia nitrogen concentration was 0.65 ± 0.06 mg/L.
2.2 | Broodstock diets and feeding protocol

Sea bass larvae collected from the hormone-induced spawning of three tested groups of broodstock were evaluated.
The broodstock were fed for 42 days on three experimental diets as follows: (a) a basal diet (BD, commercial brood-
stock diet, 40/% crude protein, 15% lipid), (b) a Palaemon-supplemented diet (PSD, 7.5% P. elegans biomass), and
(c) an Artemia-supplemented diet (ASD, 2.5% Artemia salina biomass) (Table 1). The BD was composed of fishmeal
(17%), soybean meal (46%CP) (20%), extruded full-fat soya (10%), yellow corn (15%), wheat bran (21%), corn gluten
meal (60%CP) (9%), vitamin–mineral premix and mono-calcium phosphate (4%), and fish oil (4%). The PSD was pre-
pared by adding 7.5% of adult Palaemon biomass and excluding the same portion of the BD. Palaemon (P. elegans)
was collected from Lake El-Manzala, Northern Egyptian Coast; dried (50–60 C); ground; and incorporated into
TABLE 1 Chemical composition (g/kg dry matter) and fatty acid profiles (% of total fatty acids) of Sardine, Palaemon,
Artemia, and the experimental broodstock diets used in the present study
Tested broodstock dietsa

Chemical composition Sardine Palaemon Artemia BD PSD ASD
Dry matter (DM) 240 226.0 121.9 911.3 911.8 910.6
Crude protein (CP) 747 524.0 539.3 408.4 417.0 411.6
Ether extract 57.3 91.0 113.5 150.7 146.2 149.8
Fiber 8.3 12.0 15.8 26.9 25.8 26.6
Ash 140.5 167.0 171.0 65.1 72.7 67.7
NFEb 40.69 194.0 150.4 345.0 333.7 340.1
GE MJ/kgc 20.59 19.29 19.80 21.52 21.35 21.48
LOA (18:2n-6) 5.45 12.41 2.2 3.71 4.36 3.67
LNA (18:3n-3) 0.60 0.96 8.8 0.54 0.57 0.75
ARA (20:4n-6) 1.29 2.97 3.59 0.06 0.28 0.15
EPA (20:5n-3) 0.78 14.97 11.81 0.401 1.49 0.69
DHA (22:6n-3) 13.44 8.29 0.51 0.75 1.31 0.74
ΣSFA 44.56 22.41 29.17 67.44 64.06 66.48
ΣMUFA 24.97 31.33 45.18 26.33 26.71 26.80
ΣPUFA 23.65 44.49 24.1 5.45 8.38 5.92
Total n-3 14.68 26.62 16.17 1.69 3.56 2.09
Total n-6 5.45 17.88 7.93 3.77 4.83 3.87
n3:n6 2.69 1.49 2 0.45 0.53 0.49
Cholesterol 1.44 0.40 3.7 0.09 0.11 0.18
Astaxanthin (mg/kg) 0 15 600 0 1.13 15.0
a
BD: basal diet; PSD: palaemon-supplemented diet (BD enhanced with 7.5% Palaemon biomass); ASD: artemia-supplemen-
ted diet (BD enhanced with 2.5% Artemia biomass).
b
NFE: nitrogen-free extract calculated as follow: NFE = 100 − (crude protein + ether extract + crude fiber + ash).
c
GE: Gross energy calculated on the basis of 23.6, 39.4, and 17.2 k joule gross energy/g protein, ether extract, and NFE,
respectively (National Research Council, 1993).
powdered BD. The ASD was prepared by adding 2.5% of adult Artemia biomass and excluding the same portion of
the BD. The Artemia biomass was produced by growing low price local Artemia in the laboratory as follows: the Arte-
mia dry cysts were hydrated, decapsulated, hatched, and grown with algae (Nannochloropsis oculata), and ground soy-
bean meal until adult size (Gómez et al. 1999; El-Gamal, 2010). The yielded biomass was dried (50–60 C), ground,
and incorporated into powdered BD. The PSD and ASD were perfectly mixed, pelletized to 5 mm diameter pellets
using a laboratory pelleting machine (Moulinex Electric Mincer, ME605131 1600-W, Moulinex, Groupe SEB, France),
and dried at 50–60 C for 24 hr. The reasoning for using the aforementioned ratios of Palaemon (7.5%) and Artemia
(2.5%) are attributed to the available references (Anh, 2009; Rachmansyah, Laining, & Ahmad, 2004; Sivanandavel,
Soundarapandian, & Kannupandi, 2007). Shrimp meal can be included for up to 10% in the diets of humpback grou-
per, Cromileptes altivelis (Rachmansyah et al., 2004) and at 5% in white shrimp, Penaeus indicus (Sivanandavel et al.,
2007). Dried Artemia biomass varies based on the availability and the local price compared with the cost of fish meal.
In some countries, where the cost of Artemia biomass is 1.1–1.4 US$/kg (the same price as fish meal), it is logical to
include up to 25% in marine aquafeeds (Anh, 2009). In other countries, the price of Artemia biomass reaches 22 US
$/kg (e.g., Egypt); accordingly, 2.5% might be a reasonable start.
The broodstock were fed over a course of 6 weeks from the 10th of January 2016 to the 21st of February
2016. The feeding rate was 1% of the broodstock biomass twice a day at 8:00 a.m. and 3:00 p.m. for 7 days a week.
In addition, fresh foodstuff (sardine, Sardinella spp.) was given to the broodstock in one meal per day at 2% of the
body weight (BW). The chemical compositions of the experimental diets, which were analyzed according to the stan-
dard methods of AOAC International (1995), are presented in Table 1.
2.3 | Spawning and hatching protocol

Six mature broodstock (three females and three males from each treatment) from the respective tanks were selected
for hormonal injections. On the 19th of February 2016, 2 days before the hormonal injection (40 days after starting
the feeding protocol), samples of eggs were taken via ovary biopsy using a polyethylene cannula (E.D. 1.25 mm;
I.D. 0.86 mm) to measure the average egg diameter from three female broodstock from each treatment. The percent-
age of matured eggs [egg with diameter range of 400–600 μm (mid vitellogenic), 600–800 μm (late vitellogenic)] and
hydrated eggs (≥ 800 μm) were estimated per each female (Mayer, Shackley, & Ryland, 1988). On the same day,
blood samples were collected from a caudal vesicle of anesthetized male and female broodstock (50 mg clove oil/L)
of different groups (three females and three males treatment/1) before hormonal injection, with minimum handling
stress. Blood samples (~0.5 mL/fish) were collected using a 1-mL syringe containing 50 μL of heparin and centrifuged
(4,000 rpm, 10 min, 4 C) to obtain plasma. The plasma samples were stored at −80 C until the assays of sexual hor-
mones and biochemical parameters.
On the 21st of February 2016, the broodstock were transferred to three fiberglass circular spawning tanks
(5 tons of water; 2.3 m diameter, and 1.2 m water depth) at a sex ratio of 1 female:1 male per tank. Then, broodstock
were injected intramuscularly into the base of the lateral fin with the LHRHa hormone (des-Gly10, D-Ala6-LH-RH
ethylamide acetate salt hydrate; Sigma-Aldrich, St Louis, MO). The LHRHa dose (15 μg/kg of the BW) for females
was divided into two injections; the first dose was 5 μg/kg of BW, and the second dose was 10 μg/kg of BW with a
6-hr interval with a hormonal solution volume of 0.5 mL/kg BW, while the males were injected with a dose of 5 μg/kg
of BW. The spawning tank was fitted with a surface egg collector, a 400 μm mesh-sized nylon net, installed on the
discharge water pipe.
The spawned eggs started to be released after 2 days of hormonal injection. Then, the eggs were transferred to
incubation tanks (20 L) for 4 hr at a stocking density of 10,000 eggs per liter. The incubation tanks were filled with
filtered and sterilized seawater with the same temperature and salinity as in the spawning tank. The fertilization rate
and nonviable eggs were determined. Then, the nonfertile eggs were removed by stopping the artificial aeration
(1–2 min) and siphoning the eggs from the bottom of the tank.
The fertilized eggs were incubated in hatching tanks (2 m3 conical fiberglass) for 72 hr at 15–16 C and a gentle
aeration supply through a fine air stone with 300% daily water exchange. Every day, artificial aeration was stopped in
the hatching tank for 1–2 min, and then, the dead or nonfertilized eggs that settled on the bottom of the cone were
drained. After hatching, newly hatched larvae in each hatching tank were counted before being transferred to the lar-
val rearing tanks (LRTs). The counting technique was conducted by counting larvae present in a 100 mL subsample
(five samples each time), and the average number of larvae in a sample was multiplied by the total volume of water in
the hatching tank. The effects of the maturation-enhanced diets on the ovarian development, reproductive perfor-
mance, maternal immunity, and sex-hormonal profiles of the broodstock were evaluated (El-Gamal et al., 2018, per-
sonal communication), and the final conclusion and some important data are discussed below in this study.
2.4 | Larval rearing protocol

2.4.1 | Collecting, counting, and stocking of larvae
At the beginning of the larval rearing experiment, samples (n = 3; each of 0.2 g gross fresh weight) of yolk-sac larvae
at 3 days posthatch (3 DPH) were taken from each spawning tank to determine the average initial weight and, subse-
quently, the required quantity of larvae. Six cylindrical fiberglass tanks (each 5 m3; two replicates per treatment) were
prepared, filled with filtered seawater, equipped with artificial aeration through the air blower and light control sys-
tem, and used as LRTs. About 67,000 3 DPH larvae from three hatching tanks representing the same treatment were
stocked in each LRT, with a total of 200,000 larvae per tank at a stocking density of 40 larvae/L. A total number of
1,200,000 larvae (400,000 larvae from each of the three aforementioned broodstock groups) were collected evenly
and stocked in six LRTs. After 24 hr of larvae stocking, representative water samples (n = 10; each of 1 L) were taken
from different places of the LRT using a 1,000-mL beaker, and the actual (survived) number of larvae per tank was
recalculated. This process is aimed at checking and correcting any accidental error in the total number of larvae and
also to replace any potential mortality of larvae because of manpower handling. In the present study, the newly
hatched larvae were used to evaluate the effects of maturation diets supported with crustaceans on the offspring
performance during a 42-day period of larval rearing and, afterward (up to an age of 90 days), using the common
rearing protocol used in the hatchery.
2.5 | Larval rearing conditions

The water quality parameters measured during the larval rearing period were as follows: temperature = 18 ± 1 C;
pH = 7.9 ± 0.1; DO2 saturation > 80%; salinity = 34 ± 1 ppt; total ammonia nitrogen = 0.76 ± 0.05 mg/L;
NH4 = 0.74 ± 0.08 mg/L; NH3 = 0.016 ± 0.007 mg/L. All tanks were placed within greenhouses covered with black
plastic, and the photoperiod regime was regulated at a ratio of 16:8 (light:dark) hours using artificial light according to
Moretti, Pedini, Cittolin, and Guidastri (1999). The daily water renewal rate was raised progressively from 100% dur-
ing the first 5 days to 125% until the 15th day and then increased to 150% until the end of the experiment at
45 DPH [45 DPH].
2.6 | Larval feeding protocol

The first feeding was initiated when the mouth opening was completely formed (3 DPH). The feeding regime recom-
mended for the sea bass larvae depended mainly on the stocking density and light regime. Table 2 summarizes the
applied feeding protocol in this study based on an actual technique applied in the hatchery using algae (Nannochlor-
opsis oculata), rotifer (Brachionus plicatilis), and newly hatched Artemia franciscana, and the Artemia were enriched
with Easy DHA (product of INVE®). Artificial weaning DF (product of INVE®) with different particle sizes (μm) ranging
from 80 to 500 (μm) were introduced on Days 14, 24, and 42 and were named groups DF 80–200, DF 200–300, and
DF 300–500, respectively.
ABDEL-RAHIM ET AL.
TABLE 2 Feeding protocol of European sea bass, Dicentrarchus labrax, larvae during the 42-day experimental period
Age 0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46
Algae (1,000 cells/mL)a 200 300 300 200 150 150 100
Rotifers (unit/mL)b 8 12 20 15 10
c
Small Artemia (unit/mL) 2 3 4 6 8 8 6 4
Enriched Artemia (unit/mL)d 2 2 6 8 12 12 15 20 20 15 15 12 10 5 2
DF starter (80–200 μm) − gm/m3/daye 1 2 2 2 2 2 1
DF weaning (200–300 μm) − gm/m3/day 1 1 2 3 3 4 4 5 5 3 2
3
DF weaning (300-500 μm) − gm/m /day 1 3 4
a
Algae = (Nannochloropsis oculata).
b
Rotifer = rotifer (Brachionus plicatilis).
c
small Artemia = newly hatched Artemia (Fransiscana sp).
d
enrich Artemia = enriched Artemia with Easy DHA (product of INVE®).
e
DF = dry feed (product of INVE®) with different particle sizes (μm).
7
2.7 | Measured traits

2.7.1 | Growth performance and survival
At the beginning of the larval rearing experiment, samples (n = 3; each of 0.2 g gross weight) of newly hatched fish
larvae (3 DPH) were taken from each treatment to estimate average initial weight. At the end of the 42-day larval
rearing experiment (45 DPH), representative samples (n = 5; each of 2 g gross weight) of postlarvae were collected
from each LRT for growth performance measurement. The final weight, weight gain, average daily gain (ADG), spe-
cific growth rate (SGR), final larval length, and condition factor (CF) were calculated based on the following
equations:
Weight gain ðWG;gÞ = FW − IW
Average daily gain ðg=fish=dayÞ : ADG = FW − IW=T
Specific growth rate ðSGR;%=fish=dayÞ = 100 × ½ðLn FW Þ − ðLn IW Þ=T,
where FW is the final fish weight (g), IW is the initial fish weight (g), and T is the experimental days;
h i
Condition factor ðK valueÞ = 100 × Fish weight, g=fish length3 , cm :
On the 42nd day (45 DPH), the final number of fish fry per liter was estimated by counting the postlarvae in
10 water samples taken from different positions of LRT unit using a 1,000 mL beaker. Then, the total number of PLs
per tank was calculated based on the actual number per liter multiplied by 5,000 L. Then, the survival percentage
was estimated using the following equation:
Fish survival ð%Þ = 100 × ½final number of fish=initial number of fish:
2.8 | Larval quality

2.8.1 | Swim bladder test
Swim bladder inflation tests were conducted at 25 and 75 DPH. The fish were unfed for 6 hr before each test. Fry
reared at 34 ppt salinity were exposed to a higher salinity (70 ppt) containing an anesthetic dose of 70 ppm MS-222
(Product of Sigma-Aldrich [Now MERCK], Munich, Germany) for 1 min according to Chatain and Corrao (1992).
Good-quality fry with a functional swim bladder floated on the water surface, while those that did not have a swim
bladder remained on the bottom of the container.
2.9 | Salinity/temperature stress tests

The fish fry quality was evaluated by examining their resistance to salinity and temperature shocks following the
techniques of Dhert, Lavens, and Sorgeloos (1992). A sample of 12 postlarvae (90 DPH) from each tank was exposed
to different salinity levels of 50, 45, 40, 20, and 15 ppt for periods of 30, 60, 90, and 120 min in a 10-L water bucket.
The same temperature used in the culture medium was maintained under the salinity stress tests, and gentle aeration
was supplied. Dead fish were recorded at 30-min intervals. In addition, a sample of 12 postlarvae (90 DPH) from each
tank was exposed to different temperature levels of 35, 30, 20, 15, and 10 C for periods of 30, 60, 90, and 120 min
in a 10-L water bucket. The same salinity used in the culture medium was maintained, and gentle aeration was sup-
plied. Dead fish were recorded at 30-min intervals. The cumulative mortality index (CMI) was calculated by summing
the mortality counts noted at each time interval:
CMI = Nx1 + Nx2 + Nx3 + Nx4,
where N is the number of dead fishes at times x1, x2, x3, and x4, which are 30, 60, 90, and 120 min, respectively.
2.10 | Lysozyme activity determination

2.10.1 | Sample collection and homogenization
The lysozyme activity was measured in the female broodstock, eggs, and larvae at different ages to determine the
vertical transmission of maternal immunity to offspring. Blood samples were collected without anticoagulants from
three anesthetized (using 50 mg clove oil/L) broodstock from each treatment before hormonal injection and centri-
fuged at 3,000 rpm for 10 min to obtain the serum. A serum sample was maintained at −20 C for lysozyme determi-
nation as the innate nonspecific immune parameters. A sample of fertilized eggs was harvested and used for a
lysozyme assay as the maternal immune-inherited substance. The later samples were collected at different larval ages
of 17, 31, and 45 DPH. Samples (eggs and larvae) were washed using sterile, chilled saline, after which they were fro-
zen at −20 C until the homogenization process. Frozen specimens were minced and homogenized in five volumes
(w/v) in an ice-cold sucrose buffer (0.25 M) using a Wise Tis® HG-15D Homogenizer (DAIHAN-SCIENTIFIC, India).
The homogenate was centrifuged at 10,000 rpm for 20 min at 4 C. The resultant supernatant of the tissues was col-
lected and stored at −20 C until the lysozyme assay.
2.11 | Lysozyme measurement

The lysozyme activities in the serum and tissue homogenate were measured by agarose gel cell lysis assay according
to the method described by Schultz (1987). Briefly, the lysoplates were prepared by dissolving 0.01% agarose in
0.0067 M phosphate-buffered saline (pH 6.3) at 10 C. When the agarose temperature reached 60–70 C, 500 mg of
a uniform suspension of Micrococcus lysodeikticus in 5 mL of saline was added to 1 L of agarose and mixed well, after
which the plates were poured. Exactly 25 μL of each serum sample and standard lysozyme solution were poured
carefully into each plate, which were then incubated at 28–30 C for 12–18 hr. The ring diameters of the clear zones
were measured. The diameters of the clear zones in the samples were plotted against the standard to obtain the lyso-
zyme concentration. The protein concentrations were determined according to the method of Lowry, Rosebrough,
Farr, and Randall (1951) to calculate the lysozyme-specific activity (LSA).
2.12 | Economic analysis and profitability

After completing the experiment and selling fish fry, a comprehensive economic assessment was carried out based
on the results of the current study (Part 2: larval rearing), as well as the results of the first part (reproductive perfor-
mance; carried out by the same research team). For the economic analysis of the LRT unit, the assessment was per-
formed based on two scenarios, as follows:
Scenario 1: The research team assumed that the hatchery management could achieve two successful production
cycles of sea bass offspring in a year with the same production data collected from the present study. This is a realis-
tic scenario because of the weather conditions in Egypt.
Scenario 2: The research team assumed that the hatchery management could achieve three successful production
cycles of sea bass offspring in a year with the same production data. This scenario could be easily applied using water
chillers for an earlier third production cycle or even without a chiller.
The economic analysis of this study was performed depending on the official financial data of the hatchery and
the financial data mentioned in the MSc thesis (El-Daly, 2017) for the same fish hatchery during the same year. The
annual cost components (investment, capital, and shared variables items) were distributed equally to the hatchery
tanks, including the experimental ones that shared those costs. In addition, the actual costs of the tested maturation
feed and the quantity of broodstock used to produce 200,000 newly hatched larvae stocked in each LRT were calcu-
lated for each experimental LRT unit. The evaluated economic data were: total variable costs, total production costs,
total revenue, gross margin, and net profit per LRT and/or per a thousand fish fry. The financial performance of the
current study was assessed by means of benefit–cost ratio (BCR), internal rate of return (IRR), payback period (PP),
and equilibrium point (EP) (quantitative and financial). These data are calculated using the equations listed below in
Tables 8 and 9. Data were changed from Egyptian Pound (E£) to U.S. Dollar ($) with an average exchange rate of
7.83 E£ for 1 US$.
2.13 | Statistical analyses

The results are presented as the mean ± SE for each evaluated parameter. The results were analyzed by one-way
ANOVA (SPSS, IBM Corp. Released 2013. IBM SPSS Statistics for Windows, Version 21.0. Armonk, NY: IBM Corp)
to determine the overall effects of the treatments. Duncan's multiple range test was used to discern significant differ-
ences among the treatments (larval groups) at p ± ≤ 0.05.
3 | RESULTS
3.1 | Swim bladder and survival percentage

A reasonable correlation can be observed between the tested treatments and the ratio of fish having a swim bladder
at 25 and 75 DPH in Table 3. The highest swim bladder value was significant (p ≤ 0.01), and it was recorded in the
group fed ASD. An improvement in the ratio of the swim bladder was noted at 75 DPH with a percentage ranging
between 6.5 and 7.75%. The survival (%) of postlarvae at 45 DPH was affected by the quality of the artificial feed
given to the sea bass broodstock with a significantly higher value for the group fed ASD. The increment in survival
value was 37 and 9.96% for the groups fed ASD and PSD, respectively, compared with the BD group.
3.2 | Growth performance and CF

The growth performance of the D. labrax larvae in the investigated groups during the 42-day larval rearing period of
this study is presented in Table 4. The final BW of the postlarvae spawned from the broodstock fed PSD and ASD
increased by 22.5 and 79.4%, respectively, compared with the fish in the control group (BD). The weight gains and
ADGs increased by 81.7 and 23.2% for the groups fed PSD and ASD, respectively, compared with the BD group.
Increases in the mean SGR of 6.02 and 17.3% were achieved for the groups fed PSD and ASD, respectively, com-
pared with the BD group. The length gains increased by 10.5 and 3.3% for groups fed PSD and ASD, respectively,
compared with the BD group. The CFs increased by 13.6 and 37.3% in the fish fry spawned from broodstock fed
PSD and ASD, respectively, compared with the BD treatment.
3.3 | Salinity/temperature stress tests

Table 5 illustrates the CMI of the D. labrax fry exposed to salinity stress tests at 50, 45, 40, 20, and 15 ppt for periods
of 30, 60, 90, and 120 min. The sea bass fry groups fed PSD and ASD exhibited a significant (p < 0.05) reduction in
TABLE 3 Swim bladder and survival (%) of sea bass (Dicentrarchus labrax) larvae produced from broodstock fed with
Artemia- and Palaemon biomass-supplemented diets
Treatments Swim bladder (%) at 25 DPH Swim bladder (%) at 75 DPH Survival (%)
BD 81.00 ± 1.00b 88.00 ± 1.50b 42.88 ± 0.88b
b b
PSD 84.25 ± 1.75 92.00 ± 1.00 47.15 ± 0.85b
a a
ASD 92.75 ± 0.75 99.25 ± 0.75 58.75 ± 3.75a
Level of significance 0.014 0.013 0.033
Note. Means in the same column sharing the same superscript letter are not significantly different (p > 0.05). Values are aver-
ages ± SE. ASD: Artemia-supplemented diets; BD: basal diet, PSD: palaemon-supplemented diet.
TABLE 4 Growth performance and condition factor of sea bass, Dicentrarchus labrax, postlarvae at 45 DPH
produced from broodstocks fed with Artemia- and Palaemon biomass-supplemented diets
Average daily Specific
Final weight Weight gain gain growth rate Final length Length gain Condition
Treatments (mg/pce)2 (mg/pce) (mg/pce/day) (%/day/pce) (mm/pce)3 (mm/pce) factor (CF)
BD 28.18 ± 0.40c 27.38 ± 0.40c 0.65 ± 0.01c 7.97 ± 0.04c 16.8 ± 0.2b 15.3 ± 0.2b 0.59 ± 0.03b
b b b
PSD 34.52 ± 1.19 33.72 ± 1.19 0.80 ± 0.03 8.45 ± 0.09b 17.3 ± 0.2b 15.8 ± 0.2b 0.67 ± 0.01b
ASD 50.55 ± 0.55a 49.75 ± 0.55a 1.18 ± 0.14a 9.35 ± 0.03a 18.40 ± 0.2a 16.9 ± 0.2a 0.81 ± 0.02a
Level of 0.001 0.001 0.001 0.001 0.024 0.024 0.010
significance
Note. Values are averages over the experimental period ± SE. Means in the same column sharing the same superscript letter
are not significantly different (p > 0.05). ASD: artemia-supplemented diets; BD: basal diet, PSD: palaemon-supplemen-
ted diet.
the CMI compared to the BD group at salinities of 40, 45, and 50 ppt after 120 min. The best recorded salinity toler-
ance was in favor of the ASD treatment. However, no mortalities were observed during the 20- and 15-ppt salinity
stress tests. Table 6 presents the CMI of the sea bass fry exposed to temperature stress tests at 35, 30, 20, 15, and
10 C for periods of 30, 60, 90, and 120 min. The CMI values decreased in the groups fed PSD and ASD compared
with the BD group at temperatures of 35, 30, and 10 C, and the group with the ASD treatment showed the strongest
temperature tolerance. No significant (p > 0.05) differences were detected between the treatments. The temperature
stress tests at 20 and 15 C did not cause any mortalities. The reduction ratios in the CMI of the group fed ASD were
20.8, 8.3, and 12.5% for temperatures of 10, 30, and 35 C, respectively, compared with the BD group after an expo-
sure time of 120 min. There were no significant differences between the control and the experimental treatments in
terms of temperature stress tests.
3.4 | Lysozyme-specific activity

The serum LSA of the sea bass broodstock fed PSD and ASD increased by 11.8 and 45.6%, respectively, relative to
the BD group (Table 7). A substantial increment in the LSA measured in the group fed with ASD emphasizes the
essential role of broodstock nutrition with ASD on the innate immune response during the exhausting period of the
spawning season. The peak activities of lysozyme in the fertilized eggs spawned from broodstock fed ASD and PSD
increased by 29.0 and 43.0%, respectively. Similarly, the superiority of ASD in terms of the LSA was essentially moni-
tored throughout the subsequent larval stages (17, 31, and 45 DPH) with LSA increases of 26.8, 13.8, and 20.9%,
respectively, compared with the BD group.
The general trend of the LSA demonstrated a conclusive relationship between the maternal lysozyme activity
and its concentration in the eggs (Figure 2). However, there was a gradual decrease in the LSA level after 17 and
31 DPH, after which there was an increase at 45 DPH. This finding reflects the dependency of eggs and early larval
stages on the immune substances inherited from female broodstock until they develop their own immune system.
Maternal nutrition influenced the onset of a self-immune response in the larvae as demonstrated at 45 DPH in the
groups fed ASD and PSD compared with the control group.
3.5 | Economic analysis and profitability ratio

Table 8 presents the data of the investment costs, capital costs, variable costs, and total production costs. The cost
of broodstock feeding protocol with dry diets during the 42-day maturation period increased only from 2.443 US$ in
the BD group to 5.341 US$ in the ASD group, around 3 US$ more in each LRT. Despite that, the present study
showed a slight difference in both total variable and total production cost per LRT unit around 5 US$ only among
treatments. This difference is 0.19 and 0.28% of the total variable cost and 0.11 and 0.165% of the total production
12
TABLE 5 A cumulative mortality index of European sea bass, Dicentrarchus labrax, fry subjected to salinity stress tests at 50, 45, and 40 ppt for periods of 30, 60, 90, and
120 min
Salinity stress test (ppt) and exposure time (min)
Salinity 50 ppt Salinity 45 ppt Salinity 40 ppt
Treatments 30 60 90 120 30 60 90 120 30 60 90 120
BD 16.7 ± 0.0 25.0 ± 0.0 29.2 ± 4.2a 37.5 ± 4.2a 16.7 ± 0.0 25.0 ± 0.0a 25.0 ± 0.0ab 33.3 ± 8.4 16.7 ± 8.4 25.0 ± 8.30 25.0 ± 8.3 33.3 ± 0.0a
PSD 12.5 ± 4.2 16.7 ± 8.4 25.0 ± 0.0ab 29.2 ± 4.2ab 8.3 ± 0.0 20.9 ± 4.2a 29.2 ± 4.2a 33.3 ± 0.0 4.2 ± 4.2 12.5 ± 4.2 16.7 ± 0.0 20.9 ± 4.2b
b b b b
ASD 8.3 ± 0.0 12.5 ± 4.2 16.7 ± 0.0 16.7 ± 0.0 8.3 ± 0.0 8.3 ± 0.0 12.5 ± 4.2 16.7 ± 0.0 4.2 ± 4.2 8.3 ± 0.0 8.30 ± 0.00 8.3 ± 0.0c
Level of significance1 0.192 0.374 0.074 0.051 - 0.033 0.82 0.144 0.354 0.220 0.190 0.012
Note. Sea bass fry were produced from broodstocks fed with Artemia- and Palaemon biomass-supplemented diets. Age of sea bass fry was 90 DPH, 12 pcs per replicate, and 3 replicates
per treatment. No mortalities were observed at the 20 and 15 ppt salinity stress tests. Means in the same column sharing the same superscript letter are not significantly different
(p > 0.05).
1
Values are averages ± SE. ASD: artemia-supplemented diets; BD: basal diet; PSD: palaemon-supplemented diet.
ABDEL-RAHIM ET AL.
ABDEL-RAHIM ET AL.
TABLE 6 A cumulative mortality index (CMI) of European sea bass, Dicentrarchus labrax, fry subjected to temperature stress tests at 35, 30, and 10 C for periods of 30, 60,
90, and 120 min
Temperature stress test ( C) and exposure time (min)
Temperature 35 C Temperature 30 C Temperature 10 C
Treatments 30 60 90 120 30 60 90 120 30 60 90 120
BD 12.5 ± 4.2 37.5 ± 4.2 50.0 ± 0.00 79.2 ± 4.2 4.2 ± 4.2 20.9 ± 4.2 25.0 ± 0.0 33.3 ± 0.0 16.7 ± 0.0 33.3 ± 0.0 33.3 ± 0.0 37.5 ± 4.2
PSD 8.4 ± 8.4 29.2 ± 4.2 50.0 ± 8.3 75.0 ± 8.3 8.3 ± 0.0 20.9 ± 4.2 20.9 ± 4.2 29.2 ± 4.2 16.7 ± 0.0 29.2 ± 4.2 29.2 ± 4.2 29.2 ± 4.2
ASD 4.2 ± 4.2 29.2 ± 4.2 45.9 ± 4.2 66.7 ± 8.4 4.2 ± 4.2 16.7 ± 8.4 20.9 ± 12.5 25.0 ± 0.0 12.5 ± 4.2 16.7 ± 8.3 16.7 ± 8.3 16.7 ± 8.3
1
Level of significance 0.650 0.384 0.829 0.534 0.650 0.852 0.650 0.192 0.465 0.221 0.221 0.182
Note. Sea bass fry were produced from broodstocks fed with Artemia- and Palaemon biomass-supplemented diets. Age of sea bass fry was 90 DPH, 12 pcs per replicate, and 3 replicates
per treatment. No mortalities were observed at the 20 and 15 C temperature stress tests. Values are averages ± SE. ASD: artemia-supplemented diets; BD: basal diet; PSD: palaemon-
supplemented diet.
1
No statistically significant differences among treatments were observed.
13
TABLE 7 Lysozyme-specific activity (μg/mg protein) of sea bass (Dicentrarchus labrax) broodstock serum, fertilized egg,
and different larval stages produced from broodstocks fed with Artemia- and Palaemon biomass-supplemented diets
Treatments Broodstock serum Fertilized eggs 17 DPH 31 DPH 45 DPH
BD 41.70 ± 0.15b 3.07 ± 0.17b 1.83 ± 0.09b 2.32 ± 0.06b 2.49 ± 0.06b
b ab b ab
PSD 46.61 ± 1.39 3.96 ± 0.40 1.97 ± 0.12 2.43 ± 0.09 2.74 ± 0.11ab
a a a a
ASD 60.70 ± 2.42 4.39 ± 0.25 2.32 ± 0.04 2.64 ± 0.10 3.01 ± 0.05a
Level of significance 0.00 0.05 0.02 0.09 0.01
Note. Means in the same column sharing the same superscript letter are not significantly different (p > 0.05). Values are
averages ± SE. ASD: artemia-supplemented diets; BD: basal diet; PSD: palaemon-supplemented diet.
150 BD PSD ASD

146
LYSOZYME SPECIFIC ACTIVITY (% OF CONTROL)
143
140
129
130 127
121
120
114
112
110
110 108
105
100 100 100 100 100

100
90
BS FE 17 DPH 31 DPH 45 DPH
FIGURE 2 Changes of lysozyme-specific activity (% of the control) of sea bass (Dicentrarchus labrax) broodstock
serum, fertilized egg, and different larval stages produced from broodstock fed with Artemia- and Palaemon biomass-
supplemented diets (ASD: artemia-supplemented diet; BD: basal diet; BS: broodstock; FE: Fertilized eggs; PSD:
palaemon-supplemented diet)
cost greater in the BD group compared with the other groups under Scenarios 1 and 2, respectively. Significant dif-
ferences (p ≤ 0.05) were detected concerning both total variable and total production cost per thousand fish fry pro-
duced per LRT unit. The total variable cost was 57.6–57.9% of the total production cost, with 0.05% lower cost in
PSD and ASD compared with the BD in both scenarios.
Maternal nutrition with crustacean diets positively influenced the profitability of the LRT unit. Table 9 presents the
values of the total revenue, gross margin, net profit per LRT and/or per 1,000 fry, and profitability. The total revenue per
LRT increased significantly (p ≤ 0.05) to 19.1 and 60.3% in PSD and ASD groups, respectively, compared with the BD group.
The net profit per 1,000 fry produced in each LRT unit increased by 47.3 and 114.8% in PSD and ASD compared with the
BD group under Scenario 1. Under Scenario 2, the rate of improvement was 23.6 and 54.7% in PSD and ASD groups,
respectively, compared with the BD group, with highly significant (p ≤ 0.05) differences among groups. The best IRR was
achieved in the ASD group with 15.6 and 29.26% compared with 5.3 and 13.8% per production run in the BD group in Sce-
narios 1 and 2, respectively. As a result, the estimated PP (per year) for the investment was significantly reduced in the
crustacean-enhanced groups (PSD and ASD) compared with the BD group in both scenarios. The PP reduced by 6.27 and
1.28 years in the ASD group compared with the BD group under Scenarios 1 and 2, respectively. A significant (p ≤ 0.05)
reduction in both quantitative and/or financial EP in favor of ASD, followed by the PSD group, can be detected.
TABLE 8 Investment, capital, variable, and total costs of larval rearing tanks stocked with newly hatched larvae
spawned from European sea bass broodstock fed with Artemia- and Palaemon biomass-supplemented maturation
diets*
Technical/economic Scenario #1*** Scenario #2***
criteria (per
production run) Unit§ BD** PSD** ASD** BD** PSD** ASD**
Technical criteria
Total weight of kg 3.25 2.94 2.70 3.25 2.94 2.70
broodstock (TWB)1
QTY of dry maturation kg 1.365 1.235 1.134 1.365 1.235 1.134
diet (QDMD)2
QTY of fresh kg 2.730 2.470 2.268 2.730 2.470 2.268
maturation diet
(QFMD)3
Total number of 1,000 fry/LRT 85.5 ± 1.5b 94.0 ± 2.0b 117.5 ± 7.5a 85.5 ± 1.5b 94.0 ± 2.0b 117.5 ± 7.5a
produced fry
(45 DPH), (TNPF) 4
Economic criteria
Investment cost (IC)5 $/LRT 38,314.2 38,314.2 38,314.2 25,542.8 25,542.8 25,542.8
Capital cost (CC)6 $/LRT 1,916 1,916 1,916 1,277 1,277 1,277
Variable costs
Cost of dry $/LRT 2.443 2.309 5.341 2.443 2.309 5.341
maturation diet
(CDMD)7
Cost of fresh $/LRT 1.743 1.577 1.448 1.743 1.577 1.448
maturation diet
(CFMD)8
Total cost of $/LRT 4.187 3.886 6.789 4.187 3.886 6.789
maturation diet
(TCMD)9
Cost of broodstock $/LRT 41.5 37.6 34.5 41.5 37.6 34.5
(CB)10
Labor (LC) $/LRT 827.6 827.6 827.6 552 552 552
Artemia, enrichment $/LRT 1,335.9 1,335.9 1,335.9 891 891 891
for Artemia and
rotifers, weaning
feed, etc. (OVC1)
Electricity, $/LRT 278.4 278.4 278.4 185.6 185.6 185.6
freshwater, fuel,
veterinary,
maintenance,
chemical, etc.
(OVC2)
Other variable costs $/LRT 124.4 124.2 124.2 83.7 83.5 83.5
(OVC3)
Total variable costs per $/LRT 2,612.0 2,607.6 2,607.4 1,758.0 1,753.6 1,753.4
tank (TVC1)11
Total variable costs per $/1,000 fry 30.5 ± 0.5a 27.7 ± 0.6a 22.2 ± 1.4b 20.6 ± 0.4a 18.7 ± 0.4a 14.9 ± 1.0b
fry (TVC2)12
Total production cost (TPC)
Total production cost $/LRT 4,528.0 4,523.6 4,523.4 3,035.0 3,030.6 3,030.4
per tank (TPC1)13
Total production cost $/1,000 fry 53.0 ± 0.9a 48.1 ± 1.0a 38.5 ± 2.5b 35.5 ± 0.6a 32.2 ± 0.7a 25.8 ± 1.7b
per fry (TPC2)14
TABLE 8 (Continued)
Technical/economic Scenario #1*** Scenario #2***

criteria (per
production run) Unit§ BD** PSD** ASD** BD** PSD** ASD**
Total variable %/LRT 57.69 57.64 57.64 57.92 57.86 57.86
costs/total
production cost
(TVC1 /TPC1)
Based on two scenarios of production.

Means in the same row sharing the same superscript letter are not significantly different (p > 0.05).
1
TWB = Total weight of broodstock (♀ and ♂) produced 200,000 newly hatched larvae that was discovered by Mona
El-Gamal (personal communication) based on the results of a previous experience.
2
QDMD = QTY of dry maturation diets consumed by broodstock (♀ and ♂) for 42, days to produce 200,000 newly hatched
larvae based on 1% daily feeding rate.
3
QFMD = QTY of fresh maturation diets consumed by broodstock (♀ and ♂) for 42, days to produce 200,000 newly hatched
larvae based on 2% daily feeding rate.
4
LRT = larval rearing tank (5 tons of water).
5
IC = investment cost of one unit of LRT (including all supporting units; live food, aeration, water pumps, etc.)
6
CC = capital cost (annual depreciation) was calculated based on 20 years' life span = investment cost/20 years.
7
CDMD = QDMD × 1.79, 1.87, and 4.71 $/kg diet fed BD, PSD, and ASD group, respectively.
8
CFMD = QFMD × 0.639 $/kg sardine.
9
TCMD = CDMD + CFMD.
10
CB = TWB × 38.31 $/kg.
11
TVC1 = TCMD + CB + LC + OVC1 + OVC2 + OVC3.
12
TVC2 = TVC1/TNPF.
13
TPC1 = TVC1 + CB.
14
TPC2 = TPC1/TNPF.
*Values are averages ± SE.
**ASD: artemia-supplemented diets; BD: basal diet; PSD: palaemon-supplemented diet.
***Scenario #1 (data were calculated based on two successful fry production runs per year); Scenario #2 (data were calcu-
lated based on three successful fry production runs per year).
§
One US$ = 7.83 Egyptian pound (E£) during February 2016.
4 | DISCUSSION
The first part of this comprehensive study that focused on evaluating the effects of crustacean-supplemented diets
on the reproductive performance and egg quality of broodstock indicated that absolute and relative fecundity
increased in PSD and ASD compared to the control. The relative fecundity of the PSD and ASD groups increased by
7.53 and 20.02%, respectively, more than fish fed on the BD. Values were 149,170, 160,400, and 179,030 eggs/kg
of female weight for the BD, PSD, and ASD groups, respectively. Moreover, an improvement of ovarian develop-
ment, sex hormones, egg diameter, the ratio of fertilized and hatched eggs, and lysozyme content could be detected
under crustacean-supplemented diets (El-Gamal, personal communication).
The present study demonstrates a substantial difference in the larval growth performance and vitality among the
investigated groups. Enhancing the quality of the diets fed to broodstock is becoming a more effective strategy that
is being used universally to develop nutritional reserves in the eggs and newly hatched larvae of sea bass. Enhancing
the nutritional conditions of larvae will not only promote the growth performance of larvae but also strengthen their
resistance to probable diseases (such as preweaning mortality, shock syndrome, and malformation) suffered during
the larval and postlarval development stages (Bromage & Roberts, 1995; Davis & Dinis, 2002).
One of the most remarkable phenomena that accompany feeble larvae is the increment in the ratio of fish with
noninflated swim bladders during the first 2 weeks of the larval stage (Nour, Zaki, Abdel-Rahim, & Mabrouk, 2004;
Trotter, Pankhurst, & Battaglene, 2004; Zaki, Nour, Abdel-Rahim, & Mabrouk, 2004). The presence of a functional,
developed swim bladder is one of the most critical aspects of the effective survival and development of larvae as it
gives many bony fishes the capability to control their buoyancy, sustain their depth without losing excess energy,
and chase prey efficiently. In the present study, the percentage of fish with a swim bladder was greater in 75 DPH
larvae than in 25 DPH larvae. This result is reasonable if one considers that a substantial proportion of feeble
TABLE 9 Economic analysis and profitability of larval rearing tanks stocked with newly hatched larvae spawned from European sea bass broodstock fed with Artemia- and
Palaemon biomass-supplemented maturation diets*
Scenario #1*** Scenario #2***
Economic criteria (per
§
production run) Unit BD** PSD** ASD** BD** PSD** ASD**
Total revenue, $
ABDEL-RAHIM ET AL.
Fry price, (FP) $/1,000 fry 76.63 83.01 89.40 76.63 83.01 89.40
Total revenue (TR)1 $/LRT 6,551.7 ± 114.9b 7,803.3 ± 166.03b 10,504.5 ± 670.5a 6,551.7 ± 114.9b 7,803.3 ± 166.0b 10,504.5 ± 670.5a
2 b b a b b
Gross margin (TR-TVC1) (GM) $/LRT 3,939.7 ± 114.9 5,195.7 ± 166.0 7,897.1 ± 670.5 4,793.7 ± 114.9 6,049.7 ± 166.0 8,751.1 ± 670.5a
Net profit, $
Net profit per LRT (NPPT)3 $/LRT 2023.7 ± 114.9b 3,279.7 ± 166.0b 5,981.1 ± 670.5a 3,516.7 ± 114.9b 4,772.7 ± 166.0b 7,474.1 ± 670.5a
Net profit per 1,000 fry (NPPF)4 $/1,000 fry 23.7 ± 0.9c 34.9 ± 1.0b 50.9 ± 2.5a 41.1 ± 0.6c 50.8 ± 0.6b 63.6 ± 1.7a
5 b b a b b
Benefit–cost ratio (BCR) % 0.447 ± 0.025 0.725 ± 0.037 1.322 ± 0.148 1.159 ± 0.038 1.575 ± 0.055 2.466 ± 0.221a
b a b
Profitability per tank compared % of BD 0.00 162.1 ± 8.2 295.6 ± 33.1 0.00 135.7 ± 4.7 212.5 ± 19.1a
with BD (PRT)6
Profitability per 1,000 fry % of BD 0.00 147.4 ± 4.3b 215.1 ± 10.4a 0.00 123.4 ± 1.7b 154.7 ± 4.0a
compared with BD (PRF)7
Internal rate of return (IRR)8 % 5.3 ± 0.3b 8.6 ± 0.4b 15.6 ± 1.8a 13.8 ± 0.5b 18.7 ± 0.7b 29.3 ± 2.6a
9 a b c a b
Payback period (PP) Year 9.47 ± 0.5 5.84 ± 0.3 3.20 ± 0.4 2.42 ± 0.1 1.78 ± 0.1 1.14 ± 0.1c
Equilibrium point (EP)
Quantitative (EPQ)10 1000fry/LRT 41.53 ± 0.5a 34.64 ± 0.4b 28.51 ± 0.6c 22.79 ± 0.2a 19.86 ± 0.1b 17.14 ± 0.2c
11 a b c a b
Financial (EPF) $/LRT 3,186.3 ± 37.1 2,877.6 ± 30.8 2,548.6 ± 54.1 1,745.3 ± 11.2 1,647.2 ± 10.2 1,532.9 ± 19.7c
Based on two scenarios of production.

Means in the same row sharing the same superscript letter are not significantly different (p > 0.05).
1
TR = TNPF × FP (76.63, 83.01, and 89.40 $/1,000 fry (45 DPH) for BD, PSD, and ASD, respectively).
2
GM = TR − TVC1.
3
NPPT = TR – TPC1.
4
NPF = net profit per 1,000 fry = NPPT/TNPF.
5
BCR = benefit–cost ratio = (NPPT/TPC1).
6
PRT = profitability (%) per tank = 100 × (NPPT for PSD or ASD group/average NPPT for BD group).
7
PRF = profitability (%) per 1,000 fry = 100 × (NPPF for PSD or ASD group/average NPPF for BD group).
8
IRR = internal rate of return (IRR) − % = 100 × (NPPT/IC).
9
Payback period (PP) – years = (IC/[NPPT × 2]) for Scenario #1 OR (IC/[NPPT × 3]) for Scenario #2.
10
EPQ = equilibrium point (quantitative – 1,000 fry) = CC/(FP − TVC2).
11
EPF = equilibrium point (financial – $) = CC/(1 − TVC1/TR).
*Values are averages ± SE.
***ASD: artemia-supplemented diets; BD: basal diet; PSD: palaemon-supplemented diet.
17
**Scenario #1 (data were calculated based on two successful fry production runs per year); Scenario #2 (data were calculated based on three successful fry production runs per year).
§
One US$ = 7.83 Egyptian pound (E£) during February 2016.
(noninflated) larvae will gradually die. Woolley and Qin (2010) declared that malformed swim bladders may decrease
the survival of marine finfish fingerlings by 5–10%, although the decrease may be 50% under stressful conditions. Vil-
lamizar, Garcia-Alazar, and Sanchez-Vazquez (2009) accounted for 74% swim bladder success for European sea bass
larvae, but the larvae without swim bladders died because they could not catch prey. A clear correlation between the
percentage of swim bladders and the survival (%) is verified in the present research, and this conclusion is in agree-
ment with the recommendations of Nour et al. (2004) and Zaki et al. (2004).
Many factors, such as paternally and maternally inherited factors (Peruzzi, Westgaard, & Chatain, 2007) and roti-
fer consumption (Zaki et al., 2004) in addition to the salinity (Nour et al., 2004), photoperiod, light intensity, air intake,
and temperature (Woolley & Qin, 2010), affect swim bladder inflation in marine fishes during their larval stages.
However, the interaction between swim bladder inflation and the quality of broodstock feeding has not been satis-
factorily investigated. Larger larvae produced from broodstock that were fed well-balanced diets (e.g., the group fed
ASD) may have an enhanced swimming speed, thereby supporting the better ability of those larger larvae to speedily
rise to the water surface and inhale air (Marty, Hinton, & Summerfelt, 1995; Trotter, Battaglene, & Pankhurst, 2003)
compared with smaller sizes (i.e., the BD group).
The survival rate is the most important criterion for evaluating the performance and economy of a marine fish
hatchery. The survival rates recorded in this study were comparable to values reported by other scientists, such as
lan (2011), who reported a 43.6% survival rate for sea bass. The present
Suzer, Kamac, Çoban, Saka, and Karacaog
research notably demonstrates that the enhanced survival of postlarvae spawned from broodstock fed ASD coincides
with increased lysozyme activity in eggs and larvae. This result refers to the pivotal role of lysozyme, which helps pro-
mote the development of a defense system in embryos and fish against exotic pathogens (Wang & Zhang, 2010).
Astaxanthin may also perform a prominent role in the quality of newly hatched larvae through a variety of func-
tions, including protection against severe stress conditions and antioxidant functions (Fernández-Palacios et al., 2011)
in addition to increases in vitamins A, C, and E in the body tissues of fishes. In the ongoing experiment, the amount of
astaxanthin in the ASD was higher than those in the PSD and BD. This could be another reason for the greater survival
percentage recorded for the group fed ASD. In this regard, Furuita et al. (2003) mentioned that feeding broodstock of
the Pacific sole (P. olivaceus) with vitamin A increased their fecundity, viable number of eggs, and percentage of healthy
larvae. Santiago and Gonzalo (2000) pointed out that vitamins A and C raised the survival of 3-day-old larvae produced
from largehead carp (A. nobilis) broodstock. Furthermore, the combined increase in HUFA and carotenoids significantly
boosted the larval survival rate of Sparus aurata (Scabini, Fernandez-Palacios, & Izquierdo, 2006).
Marine aquatic animals such as Artemia and Palaemon have a rich percentage of HUFA, which distinguishes them
from plant foods. The greater composition of HUFA in the ASD may be another reason for their greater survival per-
centage in the current study. This result is concurrent with the findings of Abrehouch et al. (2010), who confirmed
that common porgy (Pagrus pagrus) broodstock that were fed fresh food consisting of shrimp, squid, and Boops boops
exhibited considerably better survival percentages with regard to 2-day embryos compared with broodstock fed with
an artificial diet and sardines. The authors attribute this result to the proportion of n-3 HUFA in fish eggs, which
should not be used as a sole criterion for assessing the quality of eggs. Using Artemia enriched with HUFA and vita-
min C in the broodstock diet improved the fecundity and larval survival in guppy (Dhert et al., 2004) and angelfish
(Ershad et al., 2009). However, Parma et al. (2014) found a reasonable correlation between monounsaturated fatty
acids (MUFA), eicosopentaenoic acid (EPA), and n-6/n-3 HUFA and the survival rate of common sole larvae, while a
negative correlation was detected between DHA, n-3 PUFA, and DHA/EPA and the survival percentage. The previ-
ous authors pointed out that both low and high contents of n-3 HUFA, n-6 HUFA, and ARA in the broodstock diets
might have an adverse effect on the larval quality of the common sole. Similar to the previous work, Liang, Lu, Qian,
Zheng, and Wang (2014) reported that an excessively high DHA level in eggs has a negative effect on larval quality in
the flatfish tongue sole (Cynoglossus semilaevis).
The growth performance data obtained in the present study were nearly identical to those reported by other
researchers, including Suzer, Fırat, Saka, and Karacao
glan (2007) and Suzer et al. (2011) with a maximum SGR of
9.17%/day and a maximum weight of 48.78 mg/fish for 40-day-old sea bass postlarvae. The variations among the
investigated groups of fish larvae in our experiment are attributed mainly to inherited nutritional components, which
produced larger sizes of newly hatched larvae. Healthier newly hatched larvae were correlated with a larger size and
better growth performance. The larval quality, which is maternally inherited, is mainly assessed in yolk-sac larvae.
The diminished growth rate in fish larvae is often associated with noninflated swim bladders (Woolley & Qin, 2010)
referring to a poor ability to hunt prey and a greater metabolic requirement to control its position and swim freely in
the water column. Similarly, Trotter, Pankhurst, and Hart (2001) stated that the metabolic requirements were greater
in larvae with nonfunctional swim bladders, and larvae of yellow perch (Perca flavescens) with noninflated swim blad-
ders 20–30% lower in weight than those with normal inflated ones (Czesny, Graeb, & Dettmers, 2005).
The CF is the primary criterion utilized in fishery research (Froese, 2006), and it is applied to evaluate the quality
of fish under hatchery and growing conditions in aquaculture experiments. In the present study, the CFs were
improved in the groups of fry fed crustacean-supplemented diets, and they expressed improved health conditions
compared with the control group. This could be attributed to the content of essential nutrients released from brood-
stock to their offspring. Therefore, this could contribute to a better growth performance and fewer health problems
(such as skeletal abnormalities) that may reduce the CF (Percin & Akyol, 2009).
Stress tests are applied to assess the variations in the quality of fish fry and their resistance to abnormal environ-
mental circumstances. For instance, the salinity stress test is used to evaluate the physiological conditions of fish lar-
vae. The results of the salinity stress test in the present study concur with the findings of Dhert et al. (1992), who
discovered that sea bass larvae need fatty acids to withstand stresses. Osmotic shock experienced during the salinity
stress test undoubtedly coincides with an HUFA deficiency, which leads to different mortality rates proportional to
fatty acid reserves in the newly hatched larvae. Additional fatty acids can be sourced from the Artemia feed supple-
mented with HUFA. When Atlantic salmon smolts were subjected to salinity stress tests, the fatty acid contents in
the gills increased (Iversen et al., 2005). This increased the permeability of the branchial epithelium, leading to the
influx or outflux of ions inside or outside fish tissues under hyper- or hyposomatic rearing water. Thus, larvae with a
richer content of essential nutrients (i.e., obtained from exogenous feeding or maternal inheritance), such as HUFA
and vitamins C and E, will be more protected against stressful conditions. This results from a delay in the onset of the
inevitable destruction of some vital tissues (Jalali, Hosseini, & Imanpour, 2008).
In the present study, no mortalities were observed when sea bass fry were exposed to lower levels of salinity
(25, 20, and 15 ppt), while mortalities were detected with higher salinities starting from 40 ppt. This result is in accor-
ralı, and Tarkan (2015) and Rubio et al. (2004). However, while Rubio et al. (2004) discovered
dance with Ercan, Ag
that salinity levels between 25 and 30 ppt were better for sea bass, Ercan et al. (2015) found that the best salinity
for sea bass is 18 ppt (i.e., the isosmotic point). In the current study, a rise in the cumulative mortality rate (CMD)
was observed with an increased time exposure. This result concurs with the conclusions of Ashraf, Bengtson, and
Simpson (2010) concerning the effect of an increasing exposure time.
The effects of hyper- or hypothermal stresses on the larval stages of fishes are extremely important to under-
stand because fish fry are vulnerable to entrainment and are highly sensitive to temperature fluctuations (Lamadrid-
Rose & Boehlert, 1988). The results of the present study showed a marked reduction in the mortality rate after the
temperature stress test for the group fed a crustacean-enhanced diet. The reduction in the CMI of the group fed
ASD was 20.8, 20.8, and 12.5% for temperatures of 10, 30, and 35 C, respectively, compared with the BD group
after an exposure time of 120 min. These data agree with the findings of Barker, Townsend, and Hacunda (1981)
and Mousavi-Sabet, Ghasemnezhad, and Petrescu-Mag (2013). In particular, high temperatures have had negative
impacts on the early life stages of marine fishes (Barker et al., 1981). Furthermore, Mousavi-Sabet et al. (2013)
detected that feeding 10-day-old swordtail fry with an artificial diet and Artemia increased its protection against
high-temperature (34 C) stress tests.
Maternal immunity is of paramount interest for young fishes during the early stages of life because it provides a
defense mechanism to embryos and larvae (Swain ad Nayak 2009) during periods of immune incompetence within
the life cycle (Takemura, 1996). The present results showed an improvement of the vertical transmission of immunity
from broodfish to progeny, where the increase in lysozyme activities in offspring were concomitant to the increase in
the brood fish. Moreover, these effects continue into the larval stages of immune incompetence. The group fed ASD
showed superb results. These results match the conclusions of Lall (2000), who demonstrated that the diet is the
source of the immunological synthesis of effector and communication molecules (e.g., immunoglobulins, lysozyme,
cytokines, and eicosanoids). In addition, Cecchini et al. (2000) recorded a rise in lysozyme activity in fertilized eggs,
embryos, and fasting larvae of sea bass (D. labrax) spawned from broodstock that were fed a diet enriched in vitamin
C. In addition, Hanif, Bakopoulos, and Dimitriadis (2004) proposed that humoral specific and nonspecific immune
components could be transferred from parents, especially from immunized ones, to both eggs and larvae.
The decline in lysozyme activity during the first larval stages relative to that in the fertilized eggs in the present
study is attributed to an exhausted lysozyme content during the development process. These data concur with the
findings of Abdollahi et al. (2016), who observed a decline in the larval lysozyme levels of fertilized eggs until
10 DPH compared with eggs, after which the lysozyme level increased until 70 DPH. Immunological substances pass
to larvae via deposition by transcytosis across thecal and granulosa cells during the vitellogenesis process to imma-
ture oocytes and later to eggs and yolk-sac larvae (Kanlis et al., 1995; Picchietti et al., 2006). Therefore, managing the
broodstock immunity level during vitellogenesis and oogenesis through the optimal transfer of maternal immunity is
very important for reducing mortalities at larval/postlarval stages (Swain and Nayak 2009). The existence of proteins
with antibacterial substances such as lysozyme, which is already present in the early developmental stages (Galeotti,
1995; Takemura, 1996), plays a crucial role in the formation of a defense system against bacteria during the early life
stages in which fishes have not yet developed an effective immunocompetence. In bigger fish, lysozymes exist in sev-
eral tissues and fluids throughout the body, including fish ovaries (Takemura & Takano, 1995). Lysozymes are there-
fore involved in the defense system against pathogenic bacteria via lysis of the bacterial cell wall, where it breaks
down the linkages between N-acetylmuramic acid and N-acetylglucosamine of the peptidoglycan layers in the cell
wall (Yano, 1996).
The improved lysozyme activity in the groups fed ASD and PSD in the present study may be because of the
higher content of HUFA in the broodstock diets as HUFA enhances the production of lysozymes in females and sub-
sequent egg deposition (Reis et al., 2015; Zuo et al., 2012). Maternally transferred immunity factors transferred from
mother to offspring have been proven to enhance the immunity of fish larvae. Therefore, such factors could be
applied as an optional strategy to support the immunity of fish larvae, thereby doubling their survival rate (Zhang
et al., 2013).
The financial performance of this study adds another dimension that might contribute enough to maximize the
economic profitability of marine fish hatcheries. The crustacean-supplemented maturation diets efficiently improved
not only the vitality of the offspring but also the profitability of the LRT production unit. There has been no increase
in the total variable cost and the total production cost for the three tested treatments under both scenarios. The cost
of dry maturation diet to the total variable cost varied between 0.2 and 0.3% in the present study compared with
1.15% obtained by Watanabe, Dumas, Carroll, and Resimius (2015) for black sea bass. The production cost per thou-
sand sea bass fingerlings decreased significantly for PSD and ASD groups compared with the BD, although the total
production cost for all LRTs was very close. This is attributed to the increased number of surviving fingerlings with
bigger sizes from the LRT unit stocked with newly hatched larvae produced from broodstock fed crustacean-
supplemented diets. This is in agreement with the results of other researchers who stated that maternal nutrition has
a positive effect on the survival rate (e.g., Swain and Nayak, 2009).
The ratio of variable cost to the total production costs obtained from the present study was around 57%, com-
pared with 91.2% for European sea bass (Zacchino et al., 2014), 82.5% for black sea bass (Watanabe et al., 2015),
and 77% for humpback grouper (C. altivelis) (Siar, Johnston, & Sim, 2002). This difference may be because of the
higher cost of energy, labor, and services (Zacchino et al., 2014) and lower investment cost of the small-scale (back-
yard) hatcheries (Siar et al., 2002) compared with the higher investment cost of the hatchery in which our study was
conducted. This result supports the economic viability of PSD and ASD groups. In this context, Watanabe et al.
(2015) stated that the final stocking density (survived number) per water volume was critical to the better financial
performance of the black sea bass hatchery. Furthermore, according to Wyban and Sweeney (1991), sensitivity ana-
lyses demonstrated that increasing revenue factors (stocking density, growth rate, survival, and market price) had a
greater impact on profitability than comparable reductions in cost factors. Survival rate and selling price are the vari-
ables that significantly affect profitability (Quagrainie, 2015).
Economic performance in terms of total revenue, gross margin, and net profit (or net present value, NPV)
increased significantly in PSD and ASD compared with the BD for both scenarios. This is attributed to the higher
total revenue because of the increase in survival and the size of fingerlings (which increase fish price). Quagrainie
(2015) found that increasing survival (%) from 50 to 80% increased the profit margin by 8–9%, while the increase in
the price of hybrid striped bass from 3.7 to 4.1 US$ increased the profit margin by 11–12%. With regard to Scenario
2, the economic analysis showed financial indicators higher than Scenario 1. This is consistent with the findings of
Siar et al. (2002) and Watanabe et al. (2015) who concluded that the increased number of production cycles in a year
leads to an increase in the total number of fingerlings and thus supports the profitability of marine hatcheries.
Financial performance of the current study in terms of BCR, sometimes called “CBA” with reverse of numerator
to denominator and denominator to numerator, was more profitable in PSD and ASD treatment. BCR is a widely
accepted financial indicator used to predict the effects of a project, or an investment, verifying if the investor can
obtain a benefit or not. It is used for rational and systematic decision-making in the aquaculture sector (Di Trapani,
Sgroi, Testa, & Tudisca, 2014; Poot-López, Hernández, & Gasca-Leyva, 2014). A ratio greater than one indicates that
a project is profitable, and the higher the number, the more profitable it is. BCR increased in the crustacean-
supplemented diets under both scenarios, with the highest values (1.32–2.47) for the ASD group and the lowest ones
(0.45–1.16) for the BD group. These values are compatible with Siar et al. (2002) who recorded BCRs between 1.27
and 3.09 for Humpback grouper (C. altivelis) in backyard hatcheries.
The internal rate of return (IRR) is another financial indicator that represents the discount rate at which the pro-
ject has an NPV of zero (Larson, Wild, & Chiappetta, 2002). The higher the IRR (%), the greater the profitability.
Crustacean-supplemented maturation diets substantially increased the IRR value with 2.9- and 2.1-fold (15.6–29.3%)
higher values in ASD compared with the BD (5.3–13.8%) under Scenarios 1 and 2, respectively. The current data
were greater than 6.52–10.52%/year obtained by Watanabe et al. (2015) and lower than 356%/year surveyed by
Siar et al. (2002). Higher survival rates dramatically improve the IRR values (Quagrainie, 2015). Selling four cohorts
per year instead of only two will support the economic performance (Watanabe et al., 2015). IRR increased from
12 to 356% as a result of increasing production runs of fingerlings per year (Siar et al., 2002).
PP is also used as a financial and/or risk indicator. It estimates the required years to recover the initial invest-
ment. It is not surprising from the results of this study to reach a value of PP of nearly 1 year in ASD treatment under
Scenario 2. This is attributed to the limited producers of sea bass fingerlings, leading to the shortage of fingerlings in
the Egyptian market. Our results are in line with the results of Siar et al. (2002) when they evaluated the grouper
hatcheries in Bali, Indonesia, where 7 of the 11 hatcheries exhibited PP around 1, while our results were better than
Watanabe et al. (2015) under Scenario 1. EP is the minimum production and/or income level needed to offset total
costs at a given selling price. The lower values of fingerlings or income indicate better financial status of a treatment
(de Barros & Martins, 2012). The ASD group exhibited the lowest values in terms of both financial and production
EP. Technically and economically, LRTs under Scenario 1 actually operate below ideal production capacity, especially
for the BD group. Our results were parallel with the results of Watanabe et al. (2015) where they reported that fin-
gerlings production could be increased by up to 20% without purchasing additional larval rearing equipment.
5 | C O N CL U S I O N
The industry of sea bass hatcheries is still very lucrative in many countries worldwide given the high level of profit-
ability. Based on the results of the present study (Parts 1 and 2), it is very important to note that enhancing the
efficiency and profitability of sea bass hatchery can be achieved through the nutritional improvement of broodstock
maturation diets using inexpensive local feed additives, for example, Artemia biomass. It will not only enhance the
reproductive performance but will definitely also increase the quantity and quality of the offspring. Furthermore, the
profitability of the hatchery will significantly be improved. However, in terms of financial performance, producing
and selling three or four production runs of sea bass fingerlings per year instead of only two will support the eco-
nomic performance without additional costly infrastructure. The optimal inclusion of both Palaemon and Artemia bio-
mass in the maturation diets of sea bass requires further research. Finally, as Egypt has salt production facilities of
Artemia biomass, we recommend using local Artemia biomass or Artemia cyst to produce biomass to overcome the
expensively imported Artemia and to enhance profitability.
AC KNOWLEDG EME NT S
We thank Dr. Mohamed El Araby, Eng. Mohamed Saad, and the staff of the Marine Finfish Hatchery (Kilo 21),
GAFRD, Alexandria, for their considerable assistance with the facilities and broodstocks and their invaluable technical
support during the experimental period. Furthermore, we thank Dr. Ayman Lofty and Dr. Mohamed Toutou from the
National Institute of Oceanography and Fisheries (NIOF), Alexandria, for their help during the preparation period of
the experiment. Special thanks are also given to Mrs. Ola Laurence from Springer Nature and Mrs. Nada Nabil from
Egyptian Knowledge Bank (EKB) for the language editing of the manuscript.
OR CI D
Mohamed M. Abdel-Rahim https://orcid.org/0000-0003-2527-4780
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How to cite this article: Abdel-Rahim MM, Mansour AT, Mona MH, El-Gamal MM, El Atafy MM. To what
extent can maternal inherited immunity acquired from a crustacean-enhanced diet improve the performance
and vitality of the offspring and enhance profitability of European Sea bass (Dicentrarchus labrax)? J World
Aquacult Soc. 2019;1–25. https://doi.org/10.1111/jwas.12598
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To what extent can maternal inherited immunity acquired from a crustacean-

Article in Journal of World Aquaculture Society · February 2019

Mohamed M. Abdel-Rahim Abdallah Tageldein Mansour

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To what extent can maternal inherited immunity

artemia, Dicentrarchus labrax, fry performance, lysozyme activity,

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2 | MATERIALS AND METHODS

2.1 | Broodstock conditions

FIGURE 1 Graphical design of the experiment

2.2 | Broodstock diets and feeding protocol

Tested broodstock dietsa

2.3 | Spawning and hatching protocol

2.4 | Larval rearing protocol

2.5 | Larval rearing conditions

2.6 | Larval feeding protocol

2.7 | Measured traits

Weight gain ðWG;gÞ = FW − IW

Average daily gain ðg=fish=dayÞ : ADG = FW − IW=T

Specific growth rate ðSGR;%=fish=dayÞ = 100 × ½ðLn FW Þ − ðLn IW Þ=T,

Fish survival ð%Þ = 100 × ½final number of fish=initial number of fish:

2.8 | Larval quality

2.9 | Salinity/temperature stress tests

CMI = Nx1 + Nx2 + Nx3 + Nx4,

2.10 | Lysozyme activity determination

2.11 | Lysozyme measurement

2.12 | Economic analysis and profitability

2.13 | Statistical analyses

3.1 | Swim bladder and survival percentage

3.2 | Growth performance and CF

3.3 | Salinity/temperature stress tests

3.4 | Lysozyme-specific activity

3.5 | Economic analysis and profitability ratio

150 BD PSD ASD

100 100 100 100 100

Technical/economic Scenario #1*** Scenario #2***

Based on two scenarios of production.

Based on two scenarios of production.

Mohamed M. Abdel-Rahim https://orcid.org/0000-0003-2527-4780

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Technical/economic Scenario #1* Scenario #2*