MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
https://doi.org/10.47193/mafis.3632023010904
ABSTRACT. The aims of this research were to investigate the effects of diets with added syn-
thetic canthaxanthin (10% parafarm) and to evaluate its possible protective role under ultraviolet
radiation (UVR) in prawn Artemesia longinaris. Three isoproteic and isolipidic diets (41% protein
and 12% lipid) containing 0 (C0), 100 (C100), and 300 (C300) mg of canthaxanthin kg-1 of diet were
prepared. Before initiating the radiation experiment, prawns were fed with the different diets for a
period of 21 d in order to determine a possible accumulation of carotenoids. Afterwards, animals
were exposed to two radiation treatments for 7 d: a) photosynthetically active radiation (PAR, 400-
700 nm), and b) total radiation (PAR+UVR, 280-700 nm), under controlled conditions (19 ±2 °C,
salinity =33, pH =7). In animals exposed to PAR+UVR treatment, survival varied between 50 and
83.33% with the highest value in animals fed diet C300. At the end of the experiment, significant sta-
tistical differences were registered in integument carotenoid concentration. Under UVR stress, the
highest decrease in non-polar carotenoid and esterified astaxanthin were recorded in prawns fed
diets containing canthaxanthin. Scavenging properties were evaluated by electron resonance spec-
troscopy (EPR) using the stable 2,2-diphenyl-2-picrylhydrazyl (DPPH) radical. Prawns fed with
C300 showed the greatest activity to quench DPPH. Results suggested that dietary canthaxanthin
could be acting as an antioxidant against reactive oxygen species and produced high tolerance under
UVR stress.
Key words: Crustaceans, carotenoids, photoprotection, antioxidant activity.
Efectos de cantaxantina dietaria sobre el estres por radiacion ultravioleta en el camarón
Artemesia longinaris
RESUMEN. Los objetivos de esta investigación fueron investigar los efectos de dietas adiciona-
das con cataxantina sintética (10% parafarm) y evaluar su posible papel protector bajo la radiación
ultravioleta (RUV) en el camarón Artemesia longinaris. Se prepararon tres dietas isoprotéicas e iso-
lipídicas (41% proteína y 12% lípidos) con 0 (C0), 100 (C100) y 300 (C300) mg de cantaxantina kg-1
de dieta. Previo al experimento de radiación, los camarones fueron alimentados con las diferentes
dietas durante 21 d para determinar una posible acumulación de carotenoides. Posteriormente, los
animales fueron expuestos a dos tratamientos de radiación durante 7 d: a) radiación fotosintética-
mente activa (PAR, 400-700 nm), y b) radiación total (PAR+RUV, 280-700 nm), bajo condiciones
controladas (19 ±2 °C, salinidad =33, pH =7). En los individuos expuestos al tratamiento PAR+
RUV, la supervivencia varió entre 50 y 83,33%, con el valor más alto en animales alimentados con
dieta C300. Al final del experimento, se registraron diferencias estadísticas significativas en la con-
centración de carotenoides en el tegumento. Bajo estrés por RUV se registró la mayor disminución
233
*Correspondence:
natiarzoz@gmail.com
Received: 30 January 2023
Accepted: 10 May 2023
ISSN 2683-7595 (print)
ISSN 2683-7951 (online)
https://ojs.inidep.edu.ar
Journal of the Instituto Nacional de
Investigación y Desarrollo Pesquero
(INIDEP)
This work is licensed under a Creative
Commons Attribution-
NonCommercial-ShareAlike 4.0
International License
Marine and
Fishery Sciences
MAFIS
ORIGINAL RESEARCH
Effects of dietary canthaxanthin on ultraviolet radiation stress in prawn
Artemesia longinaris
NATALIA S. ARZOZ1, 2, *, M. ALEJANDRA MARCOVAL1, 2, A. CRISTINA DÍAZ1, 2, 3, M. LAURA ESPINO2, 3,
SUSANA M. VELURTAS1, 2 and JORGE L. FENUCCI1, 2
1Instituto de Investigaciones Marinas y Costeras (IIMyC), Universidad Nacional de Mar del Plata (UNMDP), Consejo Nacional de
Investigaciones Científicas y Técnicas (CONICET), Juan B. Justo 2550 - Mar del Plata, Argentina. 2Departamento de Ciencias Marinas,
Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata (UNMdP), Funes 3350, B7602AYL - Mar del Plata
Argentina. 3Comisión de Investigaciones Científicas, Argentina. ORCID Natalia S. Arzoz https://orcid.org/0000-0002-9034-1719
INTRODUCTION
Carotenoids are natural pigments widely dis-
tributed in nature and synthesized de novo only
by plants and some microorganisms, whereas ani-
mals depend on carotenoids supplementation
from an exogenous diet to meet their metabolic
nutritional requirements. In aquaculture, studies
on carotenoid nutrition were initially focused on
optimizing pigmentation levels (Niu et al. 2012;
Wade et al. 2015). Subsequently, after recording
the accumulation of carotenoids in tissues and
organs across different species, further studies
were conducted to explore other functions related
to the health of cultured animals, with an empha-
sis on their role as antioxidants and their effects
on growth and reproduction (Fenucci et al. 2015;
Pereira da Costa and Miranda-Filho 2020).
Astaxanthin, which is the primary pigment in
both the exoskeleton (95%) and muscle, has been
evaluated as a dietary supplement (Carreto and
Carignan 1984). Within the epidermal tissue,
astaxanthin is mainly in the monoesterified form,
while in the exoskeleton it is associated with pro-
teins forming complexes called carotenoproteins
(Wade et al. 2017). At present, alternative forms to
increase the number of carotenoids in aquatic ani-
mal production are being studied. One of them is
the addition of astaxanthin precursor carotenoids
that crustaceans can metabolize and meet their
needs (Pereira da Costa and Miranda-Filho 2020).
In recent years, global warming affected the ozone
layer increasing solar ultraviolet radiation (UVR
280-400 nm) on the Earth’s surface (Barnes et al.
2019). Different species have developed mecha-
nisms to minimize harmful effects of UVR, such
as avoiding exposure through migrations in the
water column, but not all organisms are mobile.
For these organisms, defense consists of develop-
ing protection mechanisms (Roy 2000), being the
production of structures to prevent penetration or
the production of photoprotective compounds
(PPCs) one of the most effective mechanisms
(Marcoval et al. 2020; Arzoz et al. 2022).
The most common PPCs are UV-absorbing
compounds (UACs) such as mycosporine-like
amino acids (MAAs), which act as photoprotec-
tive UV filters and present a maximum absorption
peak between 310 and 360 nm (Carreto and
Carignan 1984) and carotenoids, which absorb
within the range of 400-700 nm of visible radia-
tion (Edge 1997) and have the ability to inactivate
molecules in an electronically excited state, main-
ly those due to photosensitive reactions. Among
defense strategies, carotenoids are most likely
involved in avoiding the photooxidative damage
initiated by reactive oxygen species (Stahl and
Sies 2003). The presence of free or esterified
forms of various carotenoids has been reported in
different tissues in wild and cultured crustaceans
(Lenel et al. 1978; Castillo et al. 1982). Among
those that have been isolated from various classes
of crustaceans are non-polar carotenoids, like b-
carotene, and polar carotenoids, like astaxanthin,
equinenone, and canthaxanthin. Over the last
decades, along with the rapid growth of
aquaculture and the development of new
techniques, stressors have also emerged harming
the health of farmed animals and generating
economic losses (FAO 2020).
234 MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
de carotenoides no polares y astaxantina esterificada en camarones alimentados con dietas con cataxantina. La capacidad antioxidante
se evaluó mediante espectroscopía de resonancia electrónica (EPR) utilizando el radical estable 2,2-difenil-2-picrilhidrazilo (DPPH). Los
camarones alimentados con C300 mostraron la mayor actividad evidenciada por el decaimiento de DPPH. Los resultados sugirieron que
la cantaxantina dietaria podría estar actuando como un antioxidante contra las especies reactivas de oxígeno y producir una alta toleran-
cia bajo estrés por RUV.
Palabras clave: Crustáceos, carotenoides, fotoprotección, actividad antioxidante.
The evaluation of biological functions of
different compounds/additives, such as amino
acids, essential fatty acids, phospholipids, vita-
mins, minerals, carotenoids, different synthetic
chemical products and derivatives of bacteria,
yeasts, fungi and algae, plants and animals (Ciji
and Akhtar 2021) for stress migration, arises in
this context.
Studies to evaluate whether dietary
carotenoids, mainly astaxanthin, could improve
stress tolerance to different factors such as tem-
perature, salinity (Chen et al. 2018), nitrite (Díaz
et al. 2014) and ammonia (Pan et al. 2003) in
aquatic animals have been designed. However,
there is no information available on the possible
photoprotective role under stress by ultraviolet
radiation (UVR), which is regaining importance
for the cultivation of organisms such as peneids.
This research focused on juvenile prawn Arteme-
sia longinaris, an exclusive species found in the
South American Atlantic and distributed along
the coast from 23° S to 43° S. This species is an
important fishing resource of high commercial
value in different countries (Bauer 2020). Several
aspects of the growth and nutrition of this species
have been studied (Gimenez et al. 2002; Diaz et
al. 2017; Arzoz et al. 2022). However, the possi-
ble photoprotective role of synthetic canthaxan-
thin has not yet been evaluated and in the current
context of climate change, this is a relevant aspect
for its cultivation, which is carried out in shallow
ponds where UVR penetrates to the bottom of the
water column (Villafañe et al. 2003).
The aims of this research were to determine
the bioaccumulation of carotenoids from can-
thaxanthin-added diets in juveniles of A. longi-
naris and to evaluate their possible protective
effects under ultraviolet radiation (UVR) stress
conditions. For this purpose, the effects of the
interaction between canthaxanthin concentra-
tion in the feed and the exposure to UVR were
evaluated in terms of survival, carotenoid accu-
mulation in the integument, and free radical
scavenging.
MATERIALS AND METHODS
Experimental animals
Juveniles of A. longinaris collected with a
trawling net from Mar del Plata coastal waters
(38° 02′ S, 57° 30′ W) were transferred to J. J.
Nagera Coastal Station, National University of
Mar del Plata (38° 16′ S, 63° 30′ W). They were
kept for one week in 3,500-l cylindrical fiberglass
tanks for acclimatization to laboratory conditions
(18 ±1 °C, pH =7 and salinity = 33). Individuals
were starved for the first 24 h to empty their gut
contents and were then fed the control diet (C0)
once a day during the 10-day acclimatization
period (Table 1).
Before the radiation experiment, to determine a
possible accumulation of carotenoids, prawns
(3.09 ±0.58 g; n =90) were placed in individual
20-l glass aquaria connected to a recirculating sys-
tem having a packed-column biological filter for
the removal of nitrogenous wastes, under con-
trolled conditions (19 ±2 °C, salinity =33, pH =
7). Different dietary treatments were assigned to
30 aquaria, each aquarium containing 1 prawn
(total =90; 30 replicates per diet). Specimens
were fed three isoproteic and isolipidic diets (41%
protein and 12% lipid) for a 21-day period supple-
mented with 0 (diet C0), 100 (diet C100), and 300
(diet C300) mg of synthetic canthaxanthin kg-1 of
diet (Table 1). All ingredients were mixed and pel-
letized using the cold extrusion method (Diaz and
Fenucci 2002) and dried for 24 h at 50 °C.
At the end of the 21-day period of carotenoid
accumulation, prawns were exposed during 7 d to
two radiation treatments: photosynthetically
active radiation (PAR, range 400-700 nm) and
total radiation spectrum (PAR+UVR, range 280-
700 nm) while they continued to be fed with their
respective diets. To test each treatment, six 20-l
aquaria with two animals each were exposed to
two radiation and two feeding treatments with the
235
ARZOZ ET AL.: PHOTOPROTECTION EFFECTS IN PRAWN ARTEMESIA LONGINARIS
follow design: Treatment A: PAR-C0(control
treatment); Treatment B: PAR-C100; Treatment C:
PAR-C300; Treatment D: PAR+UVR-C0; Treat-
ment E: PAR+UVR-C100; and Treatment F:
PAR+UVR-C300. Light sources were 40 W cool-
white fluorescent bulbs (Philips) for photosyn-
thetically active radiation (PAR) and Q-Panel
UV-A-340 bulbs for UVR. Average irradiances
were 65.4 W m-2 for PAR and 20 W m-2 for UVR
(Marcoval et al. 2021), and intensities were set
according to Arzoz et al. (2022). At the end of the
exposure period, survival of different aquaria was
estimated and samples of integument and midgut
gland were taken.
Carotenoid analysis
After exposure, animals were cryoanesthezied
to take integument samples. Dissected parts were
freeze-dried and subsequently homogenized in an
argon atmosphere in darkness. Carotenoids were
analyzed following Schiedt (1993) modified by
Díaz et al. (2013). Non-polar carotenoids were
extracted three times with hexane in an argon
atmosphere in darkness. Free astaxanthin was
separated by partitioning with dimethyl sulphox-
ide (DMSO)/acetone (1:3) until colorless in an
inert atmosphere. Esterified astaxanthin was
extracted according to Napoli and Horst (1989).
Concentrations in mg g-1 tissue were calculated
using standard curves of b-carotene in hexane
(1.88 ´10-6 M), astaxanthin in DMSO/acetone
(4.19 ´10-6 M), and extinction coefficients of b-
carotene 122,000 and astaxanthin 124,000
(Perkampus 1992).
Antioxidant activity
Potential antioxidant activity in midgut gland
homogenates was determined based on the scav-
236 MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
Table 1. Composition of different diets. C0: control diet, C100: diet added with 100 mg of canthaxanthin kg-1 diet, C300: diet added
with 300 mg of canthaxanthin kg-1 diet.
Ingredient (g 100 g-1) C0 C100 C300
Fish meal (65% protein)a 48 48 48
Soybean meal (42% protein)b 17 17 17
Corn starch 20 20 20
Squid protein (85% protein) 1 1 1
Wheat bran 8.5 8.49 8.47
Canthaxanthinc 0 0.01 0.03
Fish oil 2 2 2
Fish soluble 2 2 2
Soybean lecithin 0.5 0.5 0.5
Cholesterol 0.5 0.5 0.5
Vitaminsd 0.5 0.5 0.5
aAgustinier S.A., Mar del Plata, Argentina.
bMelrico S.A., Argentina.
cSynthetic Canthaxanthin for veterinary use 10% parafarm: dark brown granulated powder with
violet reflections. Rating (11.31%, on dry basis).
dg kg-1: cholecalciferol 1.8, thiamin 8.2, riboflavin 7.8, pyridoxine 10.7, calcium panthothenate
12.5, biotin 12.5, niacin 25.0, folic acid 1.3, B12HCl 1.0, ascorbic acid (Rovimix Stay C) 39.1,
menadione 1.7, inositol 0.3, cholinechloride 0.2, a-tocopherolacetate 75, vitamin A acetate 5.0.
enging activity of the stable 2,2-diphenyl-2-
picrylhydrazyl (DPPH) free radical. Lyophilized
midgut gland (20 mg) from each treatment was
mixed with 1 ml of chloroform under an argon
atmosphere and different reaction mixtures con-
tained DPPH (6.67 ´10-6 M) were prepared
according to Diaz et al. (2014), and the radical
scavenging activity was determined using para-
magnetic electronic resonance according to Arzoz
et al. (2022).
Statistical analysis
Carotenoid concentrations were compared by
means analysis of variance (one-way ANOVA).
In addition, to compare and determine the degree
of significance of survival results, the arcsine
transformation was applied to the percentages and
one-way ANOVA was performed. Results
obtained in the tests of total antioxidant activity
were estimated by an analysis of covariance
(ANCOVA). Data were reported as mean ±stan-
dard deviation. Analyses were performed with
Vassart stats, with a significance level of a=0.05.
RESULTS
Survival percentages after a week of exposure
to different radiation and feeding treatments var-
ied between 100 and 83.33% in prawns kept
under PAR treatment, and between 50 and
83.33% for animals exposed to UVR treatment.
The highest value was observed in animals fed
diet C300, and a significant decrease (ANOVA p =
0.01) was registered in animals fed diets C0
(treatment D) and C100 (treatment E) (Figure 1).
After 28 days of feeding, significant statistical
differences were registered in carotenoid concen-
tration between treatments (Table 2). Significant
increases in non-polar carotenoid concentration
(ANOVA p =0.01 and p =0.002, respectively)
and free astaxanthin (ANOVA p =0.0008 and p =
0.03, respectively) were observed in animals
under PAR treatments (B and C), compared to
those fed diet C0(treatment A). In treatments
under UVR, animals fed diets C100 and C300
(treatments E and F) showed a statistically signif-
237
ARZOZ ET AL.: PHOTOPROTECTION EFFECTS IN PRAWN ARTEMESIA LONGINARIS
Figure 1. Survival percentages after a week of exposure to different radiation and feeding treatments. PAR: photosynthetic active
radiation, PAR+UVR: PAR +ultraviolet radiation, C0: control diet, C100, and C300: control diet supplemented with 100
and 300 mg of canthaxanthin kg-1 diet. Values are expressed as mean ±SE, n =6.
100
90
80
70
60
50
40
30
20
10
Survival (%)
PAR-C0PAR-C100 PAR-C300 PAR+UVR-C100
Treatments
83,33
100
83,33
50
66,67
83,33
0PAR+UVR-C300
PAR+UVR-C0
icant decrease in non-polar carotenoid (ANOVA
p =0.02 and p =0.01, respectively) and esterified
astaxanthin (p =0.02 and p =0.03, respectively)
compared to those of treatment D.
In all treatments, midgut gland extract had
antioxidant protective capacity evidenced by the
ability to react with the DPPH radical (Figure 2).
In the first 3 min of reaction, the percentage of
remaining DPPH was similar for all treatments
(between 82 and 95%) except for prawns fed C300
and exposed to UVR (treatment F), in which the
signal decayed drastically (60%) (ANCOVA p =
0.025). Statistically significant differences
(ANCOVA p = 0.020) were registered between
light treatments only in prawns fed diet C300, with
82% and 60% of the remaining DPPH percentage
for PAR and PAR+UVR treatments, respectively.
DISCUSSION
In aquaculture, carotenoids have been used
mainly as a source of pigment; however, potential
functions as feed additives have been inferred
from them. Several studies have demonstrated the
critical role of carotenoids in increasing stress
tolerance in aquatic animals (Lim et al. 2023).
While carotenoids from both natural and synthet-
ic sources have been evaluated, astaxanthin has
received considerable attention due to its popular-
ity as one of the most effective naturally occur-
ring antioxidants (Lim et al. 2018).
The present study represents the first assess-
ment of the effects of diets added with different
synthetic canthaxanthin concentrations on prawn
A. longinaris under UVR stress. Among peneid,
most of the studies have been carried out in
shrimp Penaeus monodon and Litopenaeus van-
namei (Wade et al. 2017). It has been demonstrat-
ed that astaxanthin improved tolerance to thermal,
osmotic (Chien et al. 2003) and ammonia stress
(Pan et al. 2003) in P. monodon. On the other
hand, Quintana López et al. (2019) observed in the
white shrimp P. vannamei that the accumulation
of astaxanthin depended on the shrimp rearing
system, with the content of esterified astaxanthin
being higher in the midgut gland, exoskeleton,
and muscles of shrimps reared under extensive
conditions compared to those reared under hyper-
intensive conditions. These results suggest that
astaxanthin could be helpful during stress condi-
tions in shrimp farming. Recently, Zhao et al.
(2022) showed that astaxanthin also improved
immunity and alleviated oxidative and ammonia
stress. Canthaxanthin is one of the intermediates
238 MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
Table 2. Carotenoids concentration (mg g-1 tissue) of juveniles A. longinaris fed diets added with different concentrations of can-
thaxanthin and exposed to different radiation treatments for a period of 21 d. PAR: photosynthetic active radiation, UVR:
ultraviolet radiation, C0: control diet, C100, C300: control diet supplemented with 100 and 300 mg of canthaxanthin kg-1 of
diet (n =3), respectively. Means in a row with different superscripts indicate significant differences (ANOVA p <0.05).
PAR PAR+UVR
A-diet C0 B-diet C100 C-diet C300 D-diet C0 E-diet C100 F-diet C300
Non-polar carotenoids 2.86 ±0.92a 5.51 ±1.01b 5.63 ±0.57b 2.83 ±0.60a 1.44 ±0.12c 1.07 ±0.38c
Free astaxanthin 1.28 ±0.43d 2.73 ±0.51e 2.78 ±0.88e 1.40 ±0.93d 0.78 ±0.11d 0.77 ±0.17d
Esterified astaxanthin 1.04 ±0.26f 1.68 ±0.49f 1.22 ±0.49f 1.42 ±0.43f 0.58 ±0.12g 0.67 ±0.25g
Total 5.18 9.92 9.63 5.65 2.8 2.51
in the metabolic pathway of carotenoids, and
shrimps can convert synthetic canthaxanthin from
feed and deposit it in their tissues as astaxanthin
(Boonyaratpalin et al. 2001). Niu et al. (2012) reg-
istered improvement in survival against low DO
stress in P. monodon fed with dietary canthaxan-
thin. Fawzy et al. (2022) evaluated the use of
diets supplemented with 0 (control), 50, 100, 200,
and 400 mg of canthaxanthin kg-1 of diet in L.
vannamei and demonstrated that supplementation
in the range of 173.73 to 202.13 mg is of primary
importance for shrimp growth and health status.
Similarly, in the present work, survival was higher
in animals fed diets added with canthaxanthin
(66.67% and 83.33%) under UVR stress. The
same percentage (83.33%) was recorded in both
treatment PAR and PAR+UVR in animals fed diet
C300. These results suggest that the inclusion of
canthaxanthin (precursor of astaxanthin) with
lower commercial cost, improved survival in A.
longinaris juveniles under UVR stress.
Quantitative and qualitative distribution of
carotenoids in aquatic animals is mainly the result
of species-specific dietary habits, metabolic
absorption, and transformation capacity of differ-
ent carotenoids. The type of pigments absorbed
and the specific absorption rates can vary consid-
erably between families or species (Meyers
2000). In the present study, animals under PAR
treatments and fed different diets exhibited sig-
nificant differences in carotenoid concentrations.
In those specimens fed diets added with canthax-
anthin, concentrations of non-polar carotenoids
and free astaxanthin were higher than in animals
fed the basal diet. Results obtained for non-polar
carotenoids coincide with previous studies in
which an increase in non-polar carotenoids, such
as b-carotene, was recorded in shrimp fed diets
added with carotenoids (Díaz et al. 2011; Pisani
et al. 2014). This difference could be due to
oxidative reactions of astaxanthin formation from
non-polar carotenoids, such as b-carotene, which
239
ARZOZ ET AL.: PHOTOPROTECTION EFFECTS IN PRAWN ARTEMESIA LONGINARIS
Figure 2. Free-radical 2,2-diphenyl-2-picrylhydrazyl (DPPH) reaction kinetics of the midgut gland of juveniles of A. longinaris
fed diets C0, C100 and C300 exposed to various light treatments over a period of 7 d. All measurements were made in
triplicate and mean values were plotted. C0: control diet; C100, C300: control diet supplemented with mg of canthaxan-
thin kg-1 diet.
100
90
80
70
60
50
40
30
20
10
DPPH remaining (%)
PAR-C0
PAR-C100
PAR-C300
PAR+UVR-C100
Time (min)
PAR+UVR-C300
PAR+UVR-C0
0 5 10 15 20
require more energy than reactions from canthax-
anthin (Ando et al. 1986). On the other hand, dif-
ferences in free astaxanthin concentration could
be due to the ability of these animals to synthe-
size astaxanthin from dietary canthaxanthin
(Wade et al. 2017). Differences in carotenoid con-
centrations were also observed in animals under
UVR treatment. A significant decrease in non-
polar carotenoid concentration and esterified
astaxanthin was recorded. These results are coin-
cident with the determination of antioxidant
activity which was higher (p =0.025, n =3) in
animals fed diets C300 compared to those fed diets
C0and C100, detecting a DPPH remnant of 60%
after 3 min.
Carotenoids play an important role in animal
health as an antioxidant through the inactivation
of free radical produced by their own metabolism
or by environmental factors (Kirti et al. 2014).
There are several mechanisms of carotenoid
action as an antioxidant, including: serving as
efficient physical quenchers of excited singlet
molecular oxygen (1O2) (Widomska et al. 2019;
Kruk et al. 2021) reacting rapidly with free radi-
cals of different origins and convert them into
more stable compounds or non-radical products
(Focsan et al. 2017, 2021); preventing the forma-
tion of free radicals through the interruption of
free radical-induced chain reactions and terminat-
ing free radical oxidations (Lai et al. 2020); and
acting as metalchelators by facilitating the con-
version of iron and copper derivatives into stable
chelate complexes (Focsan et al. 2017). Díaz et
al. (2013) demonstrated that tissues of Pleoticus
muelleri post-larvae registered (with higher
carotenoid concentrations) a higher percentage of
DPPH decay over time, probably because radi-
cals are consumed in the tissue at a rate that
depends on concentrations of protective sub-
stances. Díaz et al. (2014) registered an increase
in antioxidant capacity in post-larvae of this
species fed a diet supplemented with 100 and 300
mg of astaxanthin kg-1 of diet and exposed to
nitrite stress. In agreement with the present study,
A. longinaris under stress by UVR generated
greater antioxidant capacity when fed with a diet
added with 300 mg of canthaxanthin.
Results obtained in this study provide evidence
that the addition of synthetic canthaxanthin to the
diet of A. longinaris protects them against UVR
stress under culture conditions and suggest that
dietary canthaxanthin could be acting as an
antioxidant against reactive oxygen species gener-
ated by exposure to UVR. For the culture of A.
longinaris, the recommended concentration
should be 300 mg kg-1 of diet, amount by which
survival was not affected during exposure to UVR
and antioxidant activity was significantly higher.
These results are consistent with finding in several
fish (Rahman et al. 2016; Yi et al. 2018; Bacchetta
et al. 2019) and other crustaceans (Ettefaghdoost
et al. 2021; Fawzy et al. 2022) in which dietary
carotenoids improved their total antioxidant status
while activities of antioxidant enzymes like cata-
lase and superoxide dismutase decreased. Those
authors proposed that carotenoids are compara-
tively more potent free radical quenchers than
antioxidant enzymes and, as such, superseded
their functional importance. Although it has been
shown that carotenoid-fortified feeds stimulate the
antioxidative capacity and stress tolerance in
penaeid shrimp, more comprehensive and rigor-
ous research is still necessary to establish the
explicit functional association between antioxi-
dant defense and carotenoid-modulated immune
mechanisms and their positive contribution to
aquatic animal health and stress relief.
Competing interests
The authors declare there are no competing
interests.
Funding
This work was supported by the Consejo
Nacional de Investigaciones Científicas y Técni-
240 MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
cas (Grant Number 112201101-00019) and the
Universidad Nacional de Mar del Plata, Argentina
(Grant Number EXA15/E 827).
Data availability
Data generated or analyzed during this study
are available from the corresponding author upon
reasonable request.
Ethics
Authors confirm that the present work was
developed in compliance with the Ethical Princi-
ples for Research in Laboratory, Farm and Nature
Laboratory Animals of the National Scientific
and Technical Research Council (CONICET-
OCR-RD-20050701-1047) of Argentina.
Author contributions
Natalia S. Arzoz: writing-original draft, writ-
ing-review and editing. M. Alejandra Marcoval:
investigation, conceptualization, supervision,
writing-review and editing. A. Cristina Díaz: data
curation, formal analysis, writing-review and
editing. M. Laura Espino: software, writing-
review and editing. Susana M. Velurtas: data
curation, writing-review and editing. Jorge L.
Fenucci: project administration, funding acquisi-
tion, writing-review and editing.
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244 MARINE AND FISHERY SCIENCES 36 (3): 233-244 (2023)
