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Ciencias marinas

Print version ISSN 0185-3880

Cienc. mar vol.46 n.3 Ensenada Sep. 2020  Epub Apr 16, 2021 


Effects of frequency and feeding time on growth, food utilization, somatic indexes, and survival of juvenile white snook Centropomus viridis

Efecto de la frecuencia y la hora de alimentación sobre el crecimiento, la eficiencia alimenticia, los índices somáticos y la supervivencia en juveniles de robalo blanco Centropomus viridis

María Isabel Abdo-de la Parra1  * 

Luz Estela Rodríguez-Ibarra1 

Leonardo Ibarra-Castro1  2 

Juan Manuel Martínez-Brown1  3 

Gabriela Velasco-Blanco1 

1 Centro de Investigación en Alimentación y Desarrollo, Unidad Mazatlán, Av. Sábalo Cerritos, s/n, Estero del Yugo, 82112, Mazatlán, Sinaloa, Mexico.

2 Whitney Laboratory for Marine Bioscience, University of Florida, 9505 Oceanshore Blvd, St. Augustine, Florida 32080, USA.

3 Consejo Nacional de Ciencia y Tecnología, Av. Insurgentes Sur 1582, Col. Crédito Constructor 03940, Mexico City, Mexico.


The Pacific white snook, Centropomus viridis, is considered a species with high farming potential in Mexico due to its high economic value and overall demand in the national market. Growth in farmed fish is largely determined by the dietary regimen, which includes feeding frequency, rate, time, and cycle. The aim of the present study was to determine the feeding frequency and appropriate feeding time for farmed C. viridis juveniles in order to optimize growth and survival. Juveniles weighing 0.36 ± 0.01 g were used to evaluate 1 and up to 5 ad-libitum feeding frequencies per day, with 3 to 24-h intervals, for 6 weeks. Gained weight (GW), growth rate (GR), specific growth rate (SGR), food conversion ratio (FCR), feeding efficiency rate (FER), coefficient of variation (CV), hepatosomatic index (HI), peritoneal fat index (PFI), and survival (S) were determined for juveniles. Regardless of the feeding hours, no significant differences were found in the GR and SGR values between juveniles fed 3 times a day and those fed 5 times a day. FCR was significantly higher and FER significantly lower in treatments with juveniles fed only once a day compared with the rest of the treatments. No significant differences were found in CV, HI, and S between treatments. PFI was significantly different only between juveniles fed once and 5 times a day. Considering the results obtained in this study, we suggest feeding C. viridis juveniles 3 times a day at 6-h intervals between doses. These results will contribute to the development of biotechnology for farming this species.

Key words: feeding regimen; growth; juvenile white snooks; Centropomus viridis


El robalo blanco del pacífico, Centropomus viridis, se considera como una especie con alto potencial para su cultivo en México por su alto valor económico y gran demanda en el mercado nacional. El crecimiento de los peces en cultivo depende, en gran medida, del régimen alimenticio, el cual incluye, frecuencia, tasa, tiempo y ritmo de alimentación. El presente trabajo tuvo como objetivo determinar la frecuencia y la hora apropiada de alimentación de juveniles de C. viridis en cultivo para optimizar el crecimiento y la supervivencia. Se utilizaron juveniles de 0.36 ± 0.01 g para evaluar de 1 a 5 frecuencias de alimentación ad libitum por día, con intervalos de 3 a 24 h, durante 6 semanas. Se determinó el peso ganado (PG), la tasa de crecimiento (TC), la tasa específica de crecimiento (TEC), la tasa de conversión alimenticia (TCA), la tasa de eficiencia alimenticia (TEA), el coeficiente de variación (CV), el índice hepatosomático (IH), el índice de grasa peritoneal (IGP) y la supervivencia (S) de los juveniles. La TC y la TEC de los juveniles alimentados 3 veces al día, independientemente de las horas de alimentación, no presentaron diferencias significativas con respecto a los resultados obtenidos para los juveniles alimentados 5 veces al día. La TCA fue significativamente mayor y la TEA significativamente menor en los tratamientos con juveniles alimentados una sola vez al día en comparación con el resto de los tratamientos. No se encontraron diferencias significativas en CV, IH y S entre los tratamientos. El IGP solo presentó diferencias significativas entre los juveniles alimentados 1 y 5 veces al día. Con base en los resultados obtenidos en el presente estudio, se sugiere que los juveniles de C. viridis se alimenten 3 veces al día, con un intervalo de 6 h entre cada toma. Estos resultados ayudarán en el desarrollo de la biotecnología de cultivo de esta especie.

Palabras clave: régimen alimenticio; crecimiento; juveniles de robalo blanco; Centropomus viridis


The Pacific white snook (Centropomus viridis) has a distribution that ranges from Baja California Sur and the Gulf of California to southern Ecuador and the Galapagos Islands (Fischer et al. 1995). Species in the Centropomidae family inhabit shallow coastal waters, estuaries, rivers, and brackish lagoons and migrate to fresh waters. Juveniles and pre-adults are commonly found in mangrove areas, where they show great tolerance to salinity fluctuations given their osmoregulatory capacity (Álvarez-Lajonchère et al. 2013). Snooks are considered species with high farming potential in Mexico because of their high economic value and overall demand in the national market given the flavor, color, and texture of their meat; their high growth rates; and their ability to adapt in captivity and consume balanced feeds (Álvarez-Lajonchère and Tsuzuki 2008, Labastida-Che et al. 2013). Today, in Mexico, natural populations of snook species are overexploited (Arreguín-Sánchez and Arcos-Huitrón 2011), so biotechnological developments have begun to be made at the marine fish production pilot plant at the Research Center for Food and Development (CIAD, for its acronym in Spanish), Mazatlán (Sinaloa, Mexico), for the production of high-quality C. viridis juveniles to promote the farming of this very important species in Mexico and to develop repopulation programs (Ibarra-Castro et al. 2017).

In the marine fish farming industry, the highest operational cost is feed, so proper design of the feeding strategy is critical for a successful mariculture enterprise (Baloi et al. 2016, D’Abramo 2019). Feeding frequency affects growth, food intake, feed conversion rate and efficiency, body chemical composition, fish survival, and the quality of farming water (Biswas et al. 2010, Shipton and Hasan 2013, Costa-Bomfim et al. 2014, Rahman and Lee 2017). Juvenile marine fish need high feeding frequencies on a daily basis to attain to good performance in the culture (Schnaittacher et al. 2005, da Cunha et al. 2013, Luo et al. 2015); however, overfeeding reduces feed efficiency, increases lipid accumulation, mainly in liver and viscera, deteriorates the quality of farming water, and increases production costs (Lee and Pham 2010, Mizanur and Bai 2014, Lee et al. 2016, Guo et al. 2018). On the other hand, low feeding frequency does not provide the required nutrients for normal growth and survival of fish and, therefore, promotes size dispersion and cannibalism; furthermore, it can cause oxidative damage and immunosuppression (Tucker et al. 2006, Oh and Venmathi-Maran 2015, Tian et al. 2015). Several authors have reported optimal feeding frequencies that improve food intake, digestion, nutrient absorption, growth, and survival in juveniles marine fishes such as Limanda ferrugínea (Dwyer et al. 2002), Pseudosiaena crocea (Xie et al. 2011), Trachinotus ovatus (Wu et al. 2015), Trachinotus blochii (Hamed et al. 2016), Megalobrama amblycephala (Xu et al. 2016), Sebastes inermis (Oh et al. 2018). Likewise, it has been reported that optimal feeding frequency depends on the species, age, size, environmental factors, feed quality, and cultivation system (Hamed et al. 2016, Xu et al. 2016, Oh et al. 2018). There are currently no published data on the frequency and optimal feeding time for C. viridis; therefore, the objective of the present study was to determine the appropriate feeding frequency and time for optimal performance of C. viridis juveniles and, in this way, contribute to the development of farming biotechnology for the species.


Fish and experimental system

Centropomus viridis juveniles were obtained from the marine fish production pilot plant at CIAD in Mazatlán. The experiment was carried out in the CIAD area for bioassays, in 24 circular fiberglass tanks with black walls and white bottoms, each with 600-L capacity. The tanks had a central drain made with a 50-mm diameter PVC pipe, which was covered with a 5-mm mesh size net to prevent fish from escaping and to facilitate cleaning of the tanks. Each tank was provided with aeration and a flow-through system (approximately 6 L·min-1), with individual flow control valves to regulate flux. Seawater was pumped from Brujas Beach, Mazatlán, and passed through sand filters and cartridge filters with relative retention rating of 16 μm.

Experimental design

A completely randomized one-factor (feeding frequency) experimental design with 3 replicates was used. This design included 8 treatments, which is the number of treatments obtained after combining the number of feedings per day and feeding times (morning, noon, and evening). The assessed treatments were 1M, 1T, 2MD, 2DT, 2MT, 3MDT, 3DT, and 5MDT (Table 1). Each tank contained 20 juveniles weighing 0.36 ± 0.01 g, on average, which were fed a commercial feed for juvenile marine fish ad libitum for 6 weeks (Skretting, 0.8-1.5 mm), according to the dietary regimen (Table 1) that was randomly selected.

Table 1 Experimental design (modified from Van der Meer et al. 1997). Tested feeding frequencies and times for Centropomus viridis juveniles. 

(Treatment code)
Feeding time (h)
7:00 10:00 13:00 16:00 19:00
1T X

M = morning, T = evening, D = noon.

Experimental conditions and development

Daily food intake by juveniles and the temperature, salinity, and dissolved oxygen of water in each tank were recorded. The bottom of the tanks was siphoned daily to remove feces and food remains. During the 6 weeks of the bioassay, water temperature was kept at 29 ± 0.05 ºC, salinity at 34 ± 1.0, and dissolved oxygen at 5.5 ± 0.3 mg·L-1. To assess growth, at the end of the experiment, all juveniles from each replica were individually sedated with clove essence (0.2 mL·L-1), and after removing excess moisture with blotting paper, each individual was weighed on a digital scale with ±0.05 g precision and measured for total length (TL) with a conventional vernier. Gained weight (GW), growth rate (GR), specific growth rate (SGR), and coefficient of variation (CV) were calculated using the following formulas:

GW (g) = Average final weight (FW) - Average initial weight (IW) (1)

GR%=GWIW×100 (2)

SGR (% d-1) = 100×(ln weight at that time - ln IW)Time (3)

CV weight (%) = Standard deviation of FWFW×100 (4)

Feeding efficiency was determined using the food conversión ratio (FCR), feeding efficiency rate (FER), hepatosomatic index (HI), and peritoneal fat index (PFI):

FCR =Food intake (FI)GW (5)

FER (%) =GWFI×100 (6)

HI =Weight of liverFW×100 (7)

PFI =Wet weight of fatFw×100 (8)

The survival percentage was calculated (S):

S (%) =Final N fishInitial N fish×100 (9)

Statistical analysis of data

Percentage values were arcsine transformed to homogenize variances. All results were tested for normality (Bartlett’s test) and homoscedasticity (Levene’s test). Normal and homoscedastic data (FW, GW, GR, SGR, CV, FI, FCR, and FER) were analyzed using a one-way analysis of variance (ANOVA, P < 0.05), and significant differences between treatments were determined by Tukey’s multiple rank comparison tests (α = 0.05); data not showing a normal distribution (HI and PFI) were analyzed using a Kruskal-Wallis test (P < 0.05) and significant differences were determined using Levene’s test based on the median (Zar 1996). All statistical analyses were done using the Statgraphics Centurion XVI program v.16.204.


FW and GW were significantly higher (ANOVA, P =0.0000) in juveniles fed 5 times a day (5MDT). GR and SGR values for juveniles fed 3 times a day, regardless of feeding time (3MDT and 3DT), did not show significant differences (ANOVA, P > 0.05) with respect to the results obtained for organisms in the 5MDT treatment. Fish fed only once or twice a day, either in the morning (1M, 2MD) or in the evening (1T, 2DT), showed significantly less growth (ANOVA, P = 0.0000) than the rest of the juveniles in the other treatments. Growth in juveniles fed 2 times a day in the morning and the evening (2MT) was not significantly different (ANOVA, P > 0.05) from growth in juveniles fed 3 times a day (3MDT and 3DT). The CV showed no significant differences (ANOVA, P = 0.7548) between treatments (Table 2).

Table 2 Growth performance of juvenile Centropomus viridis during the experiment. IW, initial average weight; FW, final average weight; GW, gained weight; GR, growth rate; SGR, specific growth rate; CV, coefficient of variation. 

Treatment IW (g) FW (g) GW (g) GR (%) SGR (% d-1) CV (%)
1M 0.35 ± 0.02 2.79 ± 0.20a 2.43 ± 0.20a 685.98 ± 13.60a 4.59 ± 0.10a 31.00 ± 6.00
1T 0.34 ± 0.01 2.70 ± 0.10a 2.35 ± 0.10a 676.50 ± 66.80a 4.56 ± 0.02a 33.33 ± 4.60
2MD 0.37 ± 0.01 4.47 ± 0.30b 4.09 ± 0.30b 1064.04 ± 93.80b 5.50 ± 0.20b 34.66 ± 2.00
2DT 0.35 ± 0.01 4.49 ± 0.10b 4.13 ± 0.10b 1173.07 ± 103.40b 5.66 ± 0.10b 30.00 ± 6.00
2MT 0.36 ± 0.02 4.98 ± 0.40bc 4.61 ± 0.50bc 1247.93 ± 185.60bc 5.78 ± 0.30bc 27.33 ± 5.10
3MDT 0.36 ± 0.01 5.97 ± 0.70c 5.60 ± 0.60c 1519.45 ± 115.90cd 6.23 ± 0.10cd 32.66 ± 3.00
3DT 0.34 ± 0.02 5.65 ± 0.60bc 5.29 ± 0.60bc 1506.00 ± 173.70cd 6.19 ± 0.20cd 34.33 ± 8.10
5MDT 0.38 ± 0.01 7.32 ± 0.40d 7.00 ± 0.30d 1831.29 ± 75.47d 6.57 ± 0.10d 32.33 ± 5.50

Values (mean ± SD, n = 3) with different letters in the same column are significantly different according to Tukey’s multiple range test (( = 0.05).

Regarding feeding efficiency, juveniles in the 1M and 1T treatments showed significantly higher FCR (ANOVA, P = 0.0009) and significantly lower FER (ANOVA, P = 0.0009) in comparison with juveniles in the rest of the treatments. HI showed no significant differences (Kruskal-Wallis, P = 0.3928) between treatments. PFI was significantly different (Kruskal-Wallis, P = 0.00029) only between juveniles in the 5MDT treatment and juveniles in the 1M and 1T treatments. Regarding survival, no significant differences (P < 0.05) were observed between treatments (Table 3).

Table 3 Food utilization, somatic indexes, and survival of juvenile white snook Centropomus viridis during the experiment. FI, food intake; FCR, food conversion ratio; FER, feeding efficiency rate; HI, hepatosomatic index; PFI, peritoneal fat index; S, survival. 

Treatment FI (g) FCR FER (%) HI (%) PFI (%) S (%)
1M 3.74 ± 0.30ab 1.55 ± 0.20b 65.90 ± 12.30a 1.01 ± 0.70 2.28 ± 0.80a 100
1T 3.14 ± 0.06a 1.33 ± 0.06b 74.90 ± 3.30a 1.45 ± 0.80 2.04 ± 0.40a 100
2MD 4.49 ± 0.40bc 1.09 ± 0.07a 91.23 ± 6.90b 1.55 ± 0.10 3.25 ± 1.00ab 96.66 ± 2.80
2DT 4.19 ± 0.10abc 1.01 ± 0.05a 97.24 ± 2.40b 2.52 ± 1.90 2.96 ± 0.70ab 100
2MT 4.70 ± 0.40bc 1.02 ± 0.10a 94.98 ± 8.60b 1.53 ± 0.40 3.45 ± 0.70ab 96.66 ± 2.80
3MDT 5.82 ± 0.20d 1.04 ± 0.16a 96.70 ± 16.00b 1.50 ± 0.70 2.99 ± 1.90ab 100
3DT 5.13 ± 0.60cd 0.96 ± 0.07a 103.6 ± 8.10b 1.56 ± 0.60 3.22 ± 0.40ab 96.66 ± 2.80
5MDT 6.96 ± 0.10e 1.00 ± 0.04a 98.03 ± 1.80b 1.38 ± 0.40 4.18 ± 0.90b 100

Values (mean ± SD, n = 3) with different letters in the same column are significantly different according to Tukey’s multiple range test (( = 0.05), except for IGP, which follows Levene’s median contrast.


Several studies have shown that growth and feeding efficiency increase when feeding frequency is increased up to a certain number of times (Ribeiro et al. 2015, Baloi et al. 2016, Guo et al. 2018, Oh et al. 2019). The present study showed that growth (FCR and FER) in C. viridis juveniles increased significantly when they were fed 3 times a day and that growth was not significantly different when they were fed 5 times a day. These results are similar to those reported by Mendes-de-Oliveira et al. (2019), who tested 2 to 6 feeding frequencies per day in Centropomus undecimalis and concluded that juveniles should be fed 4 times a day for adequate growth. Moreover, Tian et al. (2015) assessed the effect of 1 to 6 feeding frequencies per day on the growth of Megalobrama amblycephala juveniles and determined that optimal feeding frequency was 3 times a day. On the contrary, if feeding frequency is reduced to 1 or 2 times a day, growth will decrease because fish cannot obtain the necessary nutrients and meet the energy requirements for their performance and somatic development; likewise, FER will decrease and FCR will increase (Salama 2008, Biswas et al. 2010, Lee and Pham 2010, Hamed et al. 2016). This was observed in this study when C. viridis juveniles fed 1 or 2 times a day showed lower GW, GR, and SGR compared with fish fed 3 and 5 times a day. FCR was significantly higher and FER was significantly lower in juveniles fed once a day. Increasing feeding frequency, on the other hand, has been reported to produce uniform fish sizes, probably because small fish have a better chance of consuming food and competition between fish in the same tank decreases (Mihelakakis et al. 2002, Tucker et al. 2006, Biswas et al. 2010, Ribeiro et al. 2015). However, in the present study, no significant differences were observed in CV values for weight between the different treatments, which has also been observed for C. undecimalis (Mendes-de-Oliveria et al. 2019) and other marine fish species such as L. ferruginea (Dwyer et al. 2002),

Pagrus auratus (Booth et al. 2008), and Platichthys flesus luscus (Küçük et al. 2014).

Regarding feeding time, the present study showed that there is no significant effect on juvenile C. viridis growth and feeding efficiency if feeding is done in the morning or the evening. This finding was also reported for Sigamus rivulatus (Barakat et al. 2011) and Colossoma macropomum (van der Meer et al. 1997). However, in some species, such as P. flesus luscus (Küçük et al. 2014) and Oplegnathus fasciatus (Oh and Venmathi-Maran 2015), reports show that food intake is higher in the first hour of the morning and in other species, such as Sardinella brasiliensis (Baloi et al. 2016) and Hapalogenys nigripinnis (Oh et al. 2019), intake is higher in the evening. In the present study, the intake of food supplied during each feeding was not evaluated, only total consumption per day, so it was not possible to determine the time of day with the highest food intake. In this study, HI in all treatments was not significantly affected by feeding frequency, and PFI was significantly higher in juveniles from the 5MDT treatment than in juveniles fed only once a day, regardless of feeding time (1M and 1T). Low PFI values suggest that during feed restriction, fish use stored fat to meet the energy requirements for growth (Baloi et al. 2016), which would explain why in the present study fish fed only once a day (1M and 1T) showed lower PFI values than fish fed 5 times a day (5MDT), since after having had insufficient food, they were possibly unable to store as much peritoneal fat as fish in the 5MDT treatment. Other studies on marine fish have also reported that PFI is affected by feeding frequency, the higher the feeding frequency, the higher the PFI values (Mizanur and Bai 2014, Baloi et al. 2016, Guo et al. 2018).

From the results obtained in the present study, it can be concluded that, under the same experimental conditions, the optimal feeding frequency for pre-fattening C. viridis juveniles is 3 times a day within 6-hour intervals. These results will contribute to the development of farming biotechnology aiming to obtain adequate growth and feeding efficiency for this species, thus enabling operational cost optimization by using only the required feed, in addition to preventing organic waste accumulation due to unconsumed food.


We thank Valerie Williams for her support in writing the English version of the abstract and correcting the manuscript, Juan Huerta for his technical assistance, and the CIAD marine fish production pilot plant staff for providing the snook juveniles. The authors state that there is no conflict of interest.

English translation by Claudia Michel-Villalobos.


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Received: January 01, 2020; Accepted: May 01, 2020

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