Aquaculture Research, 2009, 41, 53^60 doi:10.1111/j.1365-2109.2009.02302.

Activity pattern of the marine shrimp Litopenaeus

vannamei (Boone 1931) in laboratory as a function of
different feeding frequencies

Patr|¤ cia Pereira de Lima, Cibele Soares Pontes & Maria de FaŁtima Arruda
Departamento de Fisiologia, Universidade Federal do Rio Grande do Norte, Natal, RN, Brazil

Correspondence: P P de Lima, Departamento de Fisiologia, Universidade Federal do Rio Grande do Norte, Caixa Postal 1511, Campus
UniversitaŁrio, 59078-970 Natal, RN, Brazil. E-mail: pattlyma@yahoo.com.br

Abstract has increased markedly after the introduction of the

species Litopenaeus vannamei (Boone 1931) in the
Management practices on most Brazilian shrimp
1980s (Bueno 1989). Its excellent adaptation to var-
farms have resulted in high expenses for producers
ious culture conditions, especially to temperature
and damage to the environment. Applied ethology
and water salinity, has made it the main cultivated
could provide information on the pattern of shrimp
shrimp species in the country (Wainberg & Ca“mara
activity, enabling more e⁄cient feeding frequencies.
1998).
Behavioural activities of the shrimp Litopenaeus
Despite the growth in activity, management (in-
vannamei were recorded during di¡erent feeding
cluding feeding management) on most farms is still
frequencies. The shrimp were kept in aquariums
carried out empirically. The feeding frequency used,
on a 12:12 h light/dark cycle. They were fed commer-
feeding times and the number of trays per hectare
cial ration three (at 6:00, 12:00 and 18:00 hours),
are established by the producer based on traditional
four (at 6:00, 10:00, 14:00 and 18:00 hours) or
practices that can result in unnecessary expenditure
seven times per day (at 6:00. 8:00, 10:00, 12:00, 14:00,
because it is likely that not all the ration o¡ered will
16:00 and 18:00 hours). We observed animals at
be consumed. As a result, an accumulation of feed
15 min 1 aquarium 1 time periods, recording feed-
could occur, causing water and soil deterioration in
ing, substrate exploration, swimming and inactivity.
the pond (Nunes, Goddard & Gesteira 1996). This
Feeding and exploration were the highest for shrimp
could also lead to eutrophization problems in the
fed three times per day, swimming was greatest when
pond itself, as well as in adjacent areas where the ef-
animals received four feedings, whereas inactivity
£uents will be discharged.
was higher for shrimp fed seven times per day. There
Given that feeding is part of the behavioural reper-
was greater food ingestion between 12:00 and 14:00
tory of any animal species and considering the im-
hours for animals fed three and four times per day,
portance of activity patterns over a 24-h period,
whereas swimming was high mainly at 18:00 hours
understanding these aspects is critical for developing
for shrimp fed three and seven times per day. The re-
suitable management policies on shrimp farms. This
sults indicate higher forage-related activities (ex-
can be achieved through greater knowledge of
ploration/ingestion) when feed was o¡ered three or
shrimp behaviour, because ethological studies are
four times, suggesting optimization in the pattern of
now widely being used to improve the cultures of dif-
shrimp activities based on these feeding frequencies.
ferent animal species (Wechsler, Fr˛hlich, Oester,
Oswald, Troxler, Weber & Schmid 1997; Sambraus
Keywords: applied ethology, shrimp, feeding fre-
1998; Millman, Duncan, Stau¡acher & Stookey
quency, Litopenaeus vannamei
2004; Ashley 2007). Furthermore there is a need for
more studies related to the feeding frequency to be
Introduction
adopted in shrimp culture that could enable the
Shrimp culture has been gaining prominence optimization of shrimp activity. An additional advan-
throughout the world. In Brazil, commercial activity tage could be lower feed loss to the environment and

r 2009 The Authors

Journal Compilation r 2009 Blackwell Publishing Ltd 53
Feed o¡er and activities pattern in L. vannamei P P de Lima et al.
consequently greater bene¢t to the producers in
terms of sustainable culture. Greater knowledge of ac-
tivity patterns may lead to reduced arti¢cial feed costs
(ration), which has accounted for the highest produc-
tion costs of shrimp farms (Sedgwick 1979; Velasco,
Lawrence & Castille 1999; Molina, Cadena & Orellana
2000; Smith, Burford,Tabrett, Irvin & Ward 2002).
Part of the knowledge of shrimp activities has been
acquired from studies quantifying activity versus in-
activity of the animal through the use of photocells,
without discriminating between the di¡erent activ-
ities performed by these animals (Rodriguez & Naylor
1972; Hindley 1975; Reynolds & Casterlin 1979; Vance
1992; Guerao & Ribera 1996). Ethological studies ap-
plied to shrimp farming are rare (Primavera & Lebata
1995, 2000). With L. vannamei speci¢cally, some stu-
dies have been developed that show their daily activ-
ity pattern di¡erentiating each one (Pontes & Arruda
2005a, b; Pontes, Arruda, Menezes & Lima 2006).
This had led to greater knowledge of the behavioural
repertory of this species. However, many questions
related to shrimp activity remain unanswered.
One important aspect is the study of the interfer-
ence of food in inducing other shrimp activities.
According to Nunes (2000), low concentrations of or-
ganic composites in the water, such as amino acids,
sugars and nucleotides in the extracts and meals pro-
duced from ¢sh, molluscs or crustaceans when added
to arti¢cial feed, function as chemical attractants for
the shrimp stimulating foraging activity in these an-
imals. Accordingly, in addition to arti¢cial feed inges-
tion, substrate exploration, swimming and inactivity
seem to be in£uenced by the presence of food in the
surrounding environment.
A number of studies using di¡erent feeding fre-
quencies (Sedgwick 1979; Robertson, Lawrence &
Castille 1993; Josekutty & Jose 1996; Velasco et al.
1999; Smith et al. 2002; Tacon, Cody, Conquest, Diva-
karan, Forster & Decamp 2002; Carvalho & Nunes
2006) do not consider the in£uence of arti¢cial feed
on the animal’s response to this stimulus or on the
pattern of other activities. It is important to point
out that in the natural environment, feeding beha-
viour is related to the search for this resource during
the performance of other activities.
Moreover, other factors such as light intensity,
water and soil quality of the ponds, availability of nat-
ural food, time of day, larval stage and shrimp size
may in£uence arti¢cial feed consumption and conse-
quently the other activities performed by these ani-
mals (Molina-Poveda; Escobar; Gamboa-Delgado;
Cadena; Orellana & Pina 2002). Thus, the purpose of
54 Journal Compilation r 2009 Blackwell Publishing Ltd, Aquaculture Research, 41, 53^60
Aquaculture Research, 2009, 41, 53^60
this paper is to study the activity patterns of the mar-
ine shrimp L. vannamei as a function of di¡erent feed-
ing frequencies.
Material and methods
We used juvenile shrimp of L. vannamei species after 2
months of culture and a mean weight of 8.04  1.5 g
obtained from a commercial shrimp farm in the state
of Rio Grande do Norte (NE of Brazil). The tempera-
ture was maintained at 28  1 1C and salinity at
34 g L 1 throughout the experiment.
The experimental units consisted of glass aqua-
riums (0.3  0.5  0.4 m) containing 30 L of sea-
water, a closed water recirculation system, constant
aeration and continuous ¢ltration through a 3 cm
high substrate of gravel and crushed oyster shells.
Five animals were placed in each aquarium (41
shrimps m 2) although during the observations
only four of these animals were observed, owing to
the observation capacity of the researchers. In order
to recognize each animal individually during the ob-
servations, we placed di¡erent-coloured silicone
rings on the ocular peduncle. The observations began
7 days after the animals were placed in the aqua-
riums to enable them to acclimatize to the experi-
mental conditions.
The shrimp were subjected to a 12:12 hour light^
dark cycle, with behavioural observations performed
from 6:00 to 18:00 hours. Illumination in the light
phase was provided by 12 timer-controlled white
£uorescent light bulbs (32 W) distributed across the
laboratory ceiling. The last observation (18:00 hours)
was performed under individual aquarium illumina-
tion, consisting of one red incandescent light (15 W)
(Rodriguez & Naylor 1972; Hindley 1975). The white
£uorescent lights generated a mean luminosity of
180 lx for each aquarium while the red incandescent
light provided a mean luminosity of 1lx.
During the entire experiment, the shrimp were fed
an amount equivalent to 10% of the biomass of each
aquarium using pelleted shrimp ration with 35%
s
crude protein (Camaronina 35, Purina , Agribrands
do Brasil S. A., Sao Paulo, Brazil), according to the
treatment adopted: three, four or seven daily por-
tions. The same amount of ration was o¡ered inde-
pendent of the feed frequency adopted; only the
number of daily feedings varied. The ration was
o¡ered on transparent acrylic trays (4  3  2 cm),
left for 1h and removed at the end of this period.
The animals were subjected to the following feed-
ing frequencies: three feedings per day every 6 h
r 2009 The Authors
Aquaculture Research, 2009, 41, 53^60
(6:00, 12:00 and 18:00 hours); four feedings per day
every 4 h (6:00, 10:00, 14:00 and 18:00 hours); or se-
ven feedings per day every 2 h (6:00, 8:00, 10:00,
12:00, 14:00, 16:00 and 18:00 hours). Independent of
the treatment adopted, the ¢rst feeding was per-
formed immediately after the onset of the light phase
(6:00 hours) and the last after the onset of the dark
phase (18:00 hours).
The observations of each aquarium followed the
feeding regime adopted, starting immediately after
introduction of the ration and lasting 15 min, consid-
ering the following behavioural activities: (a) feeding
^ introduction of arti¢cial feed (ration) in the oral cav-
ity with subsequent ingestion; (b) exploration of the
substrate ^ continuous insertion and removal of che-
lated pereiopods with the cephalothorax inclined
slightly downward; (c) swimming ^ continuous verti-
cal or horizontal displacement or suspension in the
water column through movement of the pleopods;
and (d) inactivity ^ stationary with or without loco-
motor appendage movement. The behaviours were
recorded using the instantaneous focal method every
60 s (Martin & Bateson 2007).
A total of seven replicate tanks were applied for
each treatment; the duration of each repetition was
33 consecutive days, with four weekly observations.
The experiment included 84 animals and 490 obser-
vation hours. Given the non-adherence of the data to
normal distribution and the heterogeneity of the fac-
tor variances shown by the Shapiro^Wilks and
Levene tests, respectively, we used non-parametric
statistical analysis. To analyse behavioural activities
as a function of feeding frequency, we used the Krus-
kal^Wallis test while analysis of activities at di¡erent
feeding times was performed using the Friedman
test. For signi¢cant di¡erences, we applied the post
hoc Mann^Whitney U-test (activity versus feeding
frequency) or the Wilcoxon test (activities versus feed-
ing times). The signi¢cance level was set at 5%, and
the results in graph form were based on median va-
lues and inter-quartile range (75^25%). Di¡erent let-
ters indicate a statistically signi¢cant di¡erence.
Results
Behavioural activities as a function of feeding
frequency: feeding, substrate exploration,
swimming and inactivity
We observed signi¢cant di¡erences in feeding as a
function of the three di¡erent feeding frequencies
[Kruskal^Wallis, H (2; 8570) 5 312.043; Po0.05].
r 2009 The Authors
Journal Compilation r 2009 Blackwell Publishing Ltd, Aquaculture Research, 41, 53^60
Feed o¡er and activities pattern in L. vannamei P P de Lima et al.
The shrimp that were fed three times a day were ob-
served ingesting arti¢cial feed more frequently than
those that received four or seven feedings (Fig. 1a).
As was the case with feeding, substrate explora-
tion behaviour showed signi¢cant di¡erences
between all feeding frequencies used [Kruskal^
Wallis, H (2; 8570) 5 380.974; Po0.05], with greater
exploratory activity observed in the animals fed three
times per day (Fig. 1b).
Swimming showed di¡erences as a function of the
three feeding frequencies adopted [Kruskal^Wallis,
H (2; 8570) 5 82.314; Po0.05], the highest levels
being found in shrimp that received four daily feed-
ings (Fig.1c).
There were also di¡erences in inactivity as a func-
tion of the three feeding frequencies [Kruskal^Wallis,
H (2; 8570) 5 394.235; Po0.05], with higher inactiv-
ity recorded in the animals fed seven times a day than
those fed three or four times a day (Fig. 1d).
Behavioural activities as a function of feeding
time and di¡erent feed frequencies: feeding,
substrate exploration, swimming and
inactivity.
There were di¡erences in feeding according to feed-
ing times for the animals fed three (Friedman, w2
(2;600) 5 32.077; Po0.05), four (w2 (3;600) 5 90.774;
Po0.05) or seven times a day (w2 (6;600) 5 74.912;
Po0.05). The shrimp that received three daily feed-
ings consumed the most at 12:00 hours; those that
received four feedings ingested feed mainly at 14:00
hours. For the shrimp receiving seven daily feedings,
arti¢cial feed was ingested more frequently at 8:00 up
to 14:00 hours (Fig. 2).
Exploratory activity as a function of feeding times
showed no di¡erence for the animals fed three times
a day [Friedman, w2 (2;600) 5 1.247; P40.05]. How-
ever, there were di¡erences for the animals that re-
ceived four (w2 (3;600) 5 59.740; Po0.05) or seven
feedings (w2 (6;600) 5 86.983; Po0.05). Exploration
activity of the shrimp fed four times a day was high
at 6:00, 10:00 and 14:00 hours, decreasing only dur-
ing the last hour. The shrimp that were fed seven
times a day displayed greater exploratory activity of
the 8:00 up to 14:00 hours (Fig. 3).
The shrimp showed di¡erences in swimming when
they were fed three (Friedman, w2 (2;600) 5 20.742;
Po0.05) or seven times a day (w2 (6;600) 5 54.206;
Po0.05). When receiving three feedings, the shrimp
swam more at 12:00 and 18:00 hours. The animals
55
Feed o¡er and activities pattern in L. vannamei P P de Lima et al. Aquaculture Research, 2009, 41, 53^60

(a) (b)
Median 25%-75% Min-Max Median 25%-75% Min-Max
16 16

Substrate Exploration
13 13
(episodes/15 min.)

(episodes/15 min.)
10 10
Feeding

a b
7 7
c
4 a 4
b
c 1
1
–2 –2
3 4 7 3 4 7
Feeding frequency Feeding frequency

(c) Median 25%-75% Min-Max

(d) Median 25%-75% Min-Max
16 16
13 13 b
(episodes /15 min.)

(episodes /15 min.)

a
Swimming

10 10
Inactivity
a
7 7
b
4 a 4
c
1 1
–2 –2
3 4 7 3 4 7
Feeding frequency Feeding frequency

Figure 1 Behavioural activities: feeding (a), substrate exploration (b), swimming (c) and inactivity (d) of the shrimp
Litopenaeus vannamei (Boone 1931) as a function of feeding frequency (three, four or seven times per day). Di¡erent letters
indicate a signi¢cant di¡erence at a level of 5%.

(a) (b) (c)

Median 25%-75% Min-Max Median 25% -75% Min-Max Median 25% -75% Min-Max
16 16 16
(episodes/15 min.)

13 13 13
Feeding

10 10 10
7 b 7 7
a b
4 c 4 a a 4 a b b b
c b c d
1 1 1
–2 –2 –2
6 12 18 6 10 14 18 6 8 10 12 14 16 18
Hours Hours Hours

Figure 2 Feeding of the shrimp Litopenaeus vannamei (Boone 1931) as a function of feeding times and di¡erent frequen-
cies: (a) three times per day 5 6:00,12:00 and18:00 hours; (b) four times per day 5 6:00,10:00,14:00 and18:00 hours; and
(c) seven times per day 5 6:00, 8:00, 10:00, 12:00, 14:00, 16:00 and 18:00 hours. Di¡erent letters indicate a signi¢cant dif-
ference at a level of 5%.

that received seven daily feedings also showed more mals fed four times were more inactive at 18:00
swimming activity at 18:00 hours. There were no sig- hours. The shrimp that received seven feedings were
ni¢cant di¡erences in the animals fed four times a day more inactive at 6:00 and 16:00 hours (Fig. 5).
(w2 (3;600) 5 6.198; P40.05) (Fig. 4).
There were di¡erences in the inactivity of the ani-
mals when they received three (Friedman, w2
Discussion
(2;600) 5 34.585; Po0.05), four (w2 (3;600) 5 59.352;
Po0.05) or seven feedings (w2 (6;600) 5 75.338; The behaviours of feeding, substrate exploration,
Po0.05). When fed three times a day, the shrimp swimming and inactivity varied as a function of the
were more inactive at 6:00 and 18:00 hours; the ani- di¡erent feeding frequencies.

r 2009 The Authors

56 Journal Compilation r 2009 Blackwell Publishing Ltd, Aquaculture Research, 41, 53^60
Aquaculture Research, 2009, 41, 53^60 Feed o¡er and activities pattern in L. vannamei P P de Lima et al.

(a) (b) (c)

Substrate exploration
Median 25%-75% Min-Max Median 25% -75% Min-Max Median 25% -75% Min-Max
16 16 16

(episodes /15 min.)

13 13 13
10 10 a a a 10
b b b b b
7 7 7 c a
a
4 4 4
1 1 1
–2 –2 –2
6 12 18 6 10 14 18 6 8 10 12 14 16 18
Hours Hours Hours

Figure 3 Substrate exploration of the shrimp Litopenaeus vannamei (Boone 1931) as a function of feeding times and dif-
ferent frequencies: (a) three times per day 5 6:00, 12:00 and 18:00 hours; (b) four times per day 5 6:00, 10:00, 14:00 and
18:00 hours; and (c) seven times per day 5 6:00,8:00,10:00,12:00,14:00,16:00 and18:00 hours. Di¡erent letters indicate a
signi¢cant di¡erence at a level of 5%.

(a) (b) (c)

Median 25%-75% Min-Max Median 25% -75% Min-Max Median 25% -75% Min-Max
16 16 16
(episodes/15 min.)

13 13 13
Swimming

10 10 10
7 7 7
b b d e
4 a 4 4 a ab cd bcd ac
1 1 1
–2 –2 –2
6 12 18 6 10 14 18 6 8 10 12 14 16 18
Hours Hours Hours

Figure 4 Swimming activity of the shrimp Litopenaeus vannamei (Boone1931) as a function of feeding times and di¡erent
frequencies: (a) three times per day 5 6:00, 12:00 and 18:00 hours; (b) four times per day 5 6:00, 10:00, 14:00 and 18:00
hours; and (c) seven times per day 5 6:00, 8:00, 10:00, 12:00, 14:00, 16:00 and 18:00 hours. Di¡erent letters indicate a sig-
ni¢cant di¡erence at a level of 5%.

(a) Median 25%-75% Min-Max (b) Median 25% -75% Min-Max (c) Median 25% -75% Min-Max
16 a
16 16 b b b bd a bc
(episodes/15 min.)

13 13 c
a b 13
a a
Inactivity

10 b 10 a 10
7 7 7
4 4 4
1 1 1
–2 –2 –2
6 12 18 6 10 14 18 6 8 10 12 14 16 18
Hours Hours Hours

Figure 5 Inactivity of the shrimp Litopenaeus vannamei (Boone1931) as a function of feeding times and di¡erent frequen-
cies: (a) three times per day 5 6:00,12:00 and18:00 hours; (b) four times per day 5 6:00,10:00,14:00 and18:00 hours; and
(c) seven times per day 5 6:00, 8:00, 10:00, 12:00, 14:00, 16:00 and 18:00 hours. Di¡erent letters indicate a signi¢cant dif-
ference at a level of 5%.

The ingestion of arti¢cial feed was the highest in
the shrimp fed three times a day, indicating greater
food consumption by the animals when feeding is
more evenly spaced throughout the day. This re-
sponse could be more advantageous to the shrimp
producer in cost^bene¢t terms.
Corroborating our results, Smith et al. (2002)
assessed the e¡ect of feeding frequency on the
growth of the black tiger shrimp Penaeus monodon
(Fabricius 1798) when they were fed at di¡erent
frequencies (three, four, ¢ve or six times per day),
and found no signi¢cant di¡erences in growth, food

r 2009 The Authors

Journal Compilation r 2009 Blackwell Publishing Ltd, Aquaculture Research, 41, 53^60 57
Feed o¡er and activities pattern in L. vannamei P P de Lima et al.
conversion and animal survival. Based on these re-
sults, the authors recommend that once a nutrition-
ally suitable feed is used, frequencies of more than
three times a day provide no additional bene¢t to the
animal.
Carvalho & Nunes (2006) pointed out that the
number of feedings adopted on shrimp farms varied
as a function of the cultivation method and manage-
ment system used. These authors subjected L. vanna-
mei to di¡erent feeding frequencies (two, three, four,
¢ve or six times per day) in a grow-out pond
and found that when feed rations were adjusted
weekly only, using estimated shrimp biomass as the
criterion, more than two daily feedings were not
advantageous. Only in cases where feeding occurs
exclusively in trays, or when low nutritional and phy-
sical quality feeds are used, should multiple feedings
be considered as a strategy for reducing the food con-
version rate.
With respect to preferred feeding times at di¡erent
feeding frequencies, there was greater ingestion
between 12:00 and 14:00 hours for the animals that
received three or four daily feedings. Animals fed
seven times a day consumed more feed at 8:00 up
to 14:00 hours. This shows that feeding of shrimp, in
addition to undergoing alterations in the number of
feedings, is also in£uenced by the times at which the
feed is made available.
Molina et al. (2000) obtained similar results
when they tested L. vannamei kept in aquariums and
o¡ered arti¢cial feed twice a day at di¡erent feeding
times (8:00 and 16:00, 10:00 and 18:00, 12:00 and
20:00 or 14:00 and 22:00 hours). They found that
the shrimp fed at 12:00 or 14:00 hours consumed
more feed and had a higher ¢nal biomass, gradually
ingesting less food until 24:00 hours. They also
observed that the peak of amylase, lipase and pro-
tease enzymes occurred at 14:00 hours, independent
of feeding time. Thus, the authors suggested feeding
the shrimp larger amounts at12:00 hours, the time at
which greater food ingestion was observed and fol-
lowed by higher digestive enzyme activity.
A lower amount of arti¢cial feed was consumed at
18:00 hours independent of the feeding frequency
used (three, four or seven times per day). Because
18:00 hours was the only dark phase time recorded
in our study, our results agree with those obtained
by Pontes & Arruda (2005a). These authors o¡ered ju-
venile L. vannamei ration four times a day (8:00,
16:00, 20:00 and 4:00 hours) and found that feeding
occurred more intensely during the light phase than
it did in the dark phase.
58 Journal Compilation r 2009 Blackwell Publishing Ltd, Aquaculture Research, 41, 53^60
Aquaculture Research, 2009, 41, 53^60
Exploratory activity as well as arti¢cial feed inges-
tion showed higher values for the shrimp fed three
times a day. This may be explained by the direct rela-
tion between these two behaviours because when
searching for food in general, exploration is an im-
portant component. Exploration occurred through-
out the day and the animals fed four or seven times
daily showed a predominance of exploratory activity
during the light phase. The lowest activity was re-
corded at 18:00 hours, in contrast to the animals that
received three daily feedings, for which no di¡er-
ences were observed in relation to time.
Pontes et al. (2006), observing L. vannamei juve-
niles fed once a day at random times, showed that
substrate exploration occurred both in the light and
in the dark phase, suggesting that the search for food
occurs in both phases. The animals reached higher
peaks 7 h after the onset of the light phase, although
our data point to increased exploration up to 8 h after
the lights are turned on, after which time exploratory
activity decreases.
Swimming was more frequent among the shrimp
that received four feedings. This behaviour showed
high values, independent of the feeding frequency
adopted (three, four or seven times) at 18:00 hours,
the only observation time in the dark phase. Pontes
et al. (2006), studying the daily behavioural pattern
of L. vannamei over a 24-h period, found that they
swam predominantly in the dark phase.
With respect to inactivity, it was observed that the
shrimp were more inactive when they were fed seven
times a day. The shrimp that received three or seven
feeds remained stationary at 6:00 hours, becoming
more active over the course of the day until they once
again registered high values for this behaviour at the
end of the day. Owing to large energy expenditures
during swimming, the shrimp seem to alternate be-
tween moments of activity and inactivity. This may
be due to the relation between activity/inactivity
and oxygen consumption.
Dall (1986) related shrimp activity with oxygen
consumption in his study performed with Penaeus es-
culentus (Haswell 1879) to measure the metabolic
routine of shrimp through oxygen consumption. He
found small £uctuations in O2 consumption during
the day when the shrimp were more inactive. At
night, when they become more active, oxygen con-
sumption triples.
Primavera & Lebata (2000), based on 25 hourly
laboratory observations, also found higher activity
levels between Metapenaeus ensis (De Haan 1844),
Penaeus latisulcatus (Kishinouye1896) and Fennerope-
r 2009 The Authors
Aquaculture Research, 2009, 41, 53^60
naeus merguiensis (De Man 1888) juveniles during the
dark phase. M. ensis and P. latisulcatus juveniles dis-
played a pattern of burrowing themselves into the
substrate during the day and emerging at night. Diur-
nal burrowing was greater in Metapenaeus than in
Penaeus while activities above the substrate (clean-
ing, feeding, sheltering, inactivity, swimming and
crawling) were more frequent in Penaeus than in Me-
tapenaeus species. This pattern was less pronounced
in F. merguiensis, which is equally active in the light
and dark phases.
Thus, it can be a⁄rmed that feeding frequency
in£uenced the pattern of shrimp activities. The
highest feeding and exploration levels resulted from
three feedings per day, followed by four feedings.
When fed seven times a day, the animals explored less
and were more inactive. Given that exploration is
an activity related to the search for food, the less
frequent feedings (three or four times a day in the
light phase) seem to optimize activity patterns
directed towards both searching for and consuming
food, suggesting that the shrimp should receive three
daily feedings. Three feedings per day is bene¢cial
both to the animal and to the producer, who can
improve management and achieve better business
results.
Acknowledgments
We would like to thank the Conselho Nacional de
Desenvolvimento Cient|¤ ¢co e Tecnolo¤gico (CNPq),
Coordenacao de Aperfeicoamento de Pessoal de N|¤ vel
Superior (Capes), the Papeba shrimp farm and the
Department of Physiology of the Universidade Feder-
al do Rio Grande do Norte for their support in devel-
oping this study.
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