Aquaculture Research, 2008, 39, 276^282 doi:10.1111/j.1365-2109.2007.01894.

Antipredator responses to overhead fright stimuli in

hatchery-reared and wild European sea bass
(Dicentrarchus labrax L.) juveniles

Stefano Malavasi,Vyron Georgalas, Danilo Mainardi & Patrizia Torricelli

Dipartimento di Scienze Ambientali, UniversitaØ Ca’ Foscari di Venezia, Campo della Celestia,Venezia, Italy

Correspondence: S Malavasi, Dipartimento di Scienze Ambientali, UniversitaØ Ca’ Foscari di Venezia, Campo della Celestia, Castello 2737/b,
30122 Venezia, Italy. E-mail: mala@unive.it

Abstract It has been observed widely that hatchery-reared

In this study, shoals of hatchery-reared and wild sea
bass juveniles (Dicentrarchus labrax L.) were tested
for di¡erences in their antipredator responses to-
wards a visual (shadow) and a mechanical (dummy
bill) overhead stimulus. Two behavioural variables ^
distance from the bottom and freezing duration ^
were measured during post-stimulus phases of each
test and compared between wild and hatchery-
reared shoals, composed of 10 juveniles each. The
results showed that in both hatchery-reared and wild
juveniles, stimulus exposure elicited a signi¢cant
decrease in the mean shoal distance from the bottom.
Similarly, individuals from both groups engaged a
freezing reaction, but the mean freezing duration
was signi¢cantly higher in wild- than in hatchery-
reared juveniles. Results are discussed in the light of
their relevance for the enhancement of restocking
programmes.
Keywords: antipredator behaviour, sea bass,
restocking, reared juveniles
Introduction
Juvenile ¢sh are vulnerable to predation (Houde1987;
Webster 2002; Snickars, Sandstrom & Mattila 2004),
as the highest mortality occurs in the youngest age
classes of the population (Hempel 1963; Houde
2002). Antipredator behaviour is therefore an impor-
tant adaptive component of the behavioural reper-
toire of the early stage of ¢sh species (Giles 1984).
¢sh released into the wild experience lower levels of
survival than do wild ¢sh (Suboski & Templeton1989;
Brown & Smith 1998; Einum & Fleming 2001; Brown
& Day 2002) and this is partly due to behavioural def-
icits in antipredator behaviour incurred through
rearing in a psychosensory-deprived environment
(Howell 1994; Olla, Davis & Ryer 1994; Brown & Day
2002; Huntingford 2004; Malavasi, Georgalas, Lugli,
Torricelli & Mainardi 2004). The assessment of the
behavioural di¡erences between hatchery-reared
and wild juveniles is therefore needed as a starting
point for any restocking programme.
In the framework of assessing the biological qual-
ity of juvenile ¢sh from northern Adriatic popula-
tions of reared ¢sh species (sea bass, sea bream and
mullets), di¡erences between hatchery-reared and
wild sea bass juveniles in the behavioural responses
towards a ¢sh predator were recently analysed
(Malavasi et al. 2004).
In this paper, a further component of the antipre-
dator behaviour of sea bass juveniles was investi-
gated, i.e. the behavioural responses towards attacks
from above, as performed by a potential bird predator.
As stated by Petersson and Jarvi (2006), attacks from
above received by ¢sh in the hatchery environments
are normally not e¡ective and arti¢cial selection
may favour non-responding individuals. Moreover,
Gotceitas and Godin (1991) showed that a ¢sh
responds to the simulated overhead £ight of a bird
predator by reducing their activity.
Antipredator responses towards attacks from
above consist usually of di¡erent components, such
as £eeing (burst swimming) and freezing (motionless

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276 Journal Compilation r 2008 Blackwell Publishing Ltd
Aquaculture Research, 2008, 39, 276^282
resting on the bottom) (Giles 1984; Csanyl 1985;
Giacquinto & Volpato 2001; Brown & Dreier 2002;
Lehtiniemi 2005; Petersson & Jarvi 2006). It has been
shown that increased frequency of freezing is a reli-
able indicator of antipredator behaviour in ¢sh
species, as a response towards chemical alarms
(Giacquinto & Volpato 2001; Brown & Dreier 2002;
Vilhunen & Hirvonen 2003), ¢sh predators (Csanyl
1985; Vehanen 2003) and models of bird predators
(Giles 1984; Giles & Huntingford 1984; Gotceitas &
Godin1991). A recent paper testing for the di¡erences
between hatchery-reared and wild coho salmon in
the fright responses towards an overhead stimulus
did not ¢nd any signi¢cant di¡erence between hatch-
ery and wild fry (Ryer 2004), contrary to the expecta-
tions based on the general evidence showing
di¡erences in antipredator behaviour between wild
and hatchery ¢sh (see Huntingford 2004, for a
review).
In this paper, components of the antipredator re-
sponses (that is freezing duration and distance from
the bottom after £eeing) towards two di¡erent kinds
of overhead fright stimuli were measured and com-
pared between groups of hatchery-reared and wild
European sea bass juveniles (Dicentrarchus labrax L.).
Two experimental sets were arranged, in order to as-
sess quantitatively the antipredator responses of ¢sh
juvenile shoals towards an overhead visual stimulus
(shadow created by a £ying object above the water
surface) and an overhead mechanical stimulus
(using a dummy bill). The results were discussed in
the light of the current knowledge on the di¡erences
in antipredator behaviour between hatchery and
wild ¢sh.
Materials and methods
Experimental animals and housing conditions
Juvenile sea bass were captured at the southern coast
of Italy (Caserta, Campania, southern Italy; 41104 0 N;
14120 0 E) by professional fry catchers and transported
to Val Figheri (southern Venice Lagoon, northern
Italy; 45118 0 N;12110 0 E), where the indoor experimen-
tal and hatchery facilities were located. Juvenile sea
bass obtained in this way are routinely seeded in the
Venice Lagoon to restock the local sea bass popula-
tion, thereby sustaining the traditional form of aqua-
culture named ‘vallicoltura’ (Ardizzone, Cataudella &
Rossi 1988).
Once in the hatchery, and after 4 h of acclimatiza-
tion, a stock of 400 wild ¢sh was equally distributed
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Antipredator reaction in sea bass juveniles S Malavasi et al.
into four 225 L (100 cm
50 cm
45 cm), comple-
tely bare, glass aquarium tanks, each equipped with
a continuous water-£ow system. The side walls of
each tank were covered with thick, dark-blue-
coloured paper in order to prevent ¢sh from excessive
stress and/or conditioning. Water temperature was
measured daily and averaged 19  1 1C throughout
the duration of the experiments. Water salinity was
maintained at 20 g L 1 and the photoperiod regime
corresponded to the natural cycle for the latitude
and time of year. Fish were fed twice daily with live
food (Artemia spp.).
Hatchery-reared juvenile sea bass were obtained
from captive-induced spawns in the hatchery facil-
ities of Val Figheri. The broodstock had spent 7^8
years in captivity and originated from the Venice La-
goon. The larvae, reared following the mesocosm
technique (Shields 2001), were held in tanks of
64 m3, at a larval density of 5^6 individuals L  1.
A batch of 113-day-old juvenile sea bass captured
from the original tanks was allocated randomly to
four aquaria identical to those housing the wild
juveniles. Care was taken to use hatchery-reared
juveniles of the same average size as that of the wild
¢sh. A random sample of 30 juveniles from wild and
hatchery-reared stocks was captured, anaesthetized
(using MS 222, Sigma-Aldrich, St Louis, MO, USA), sa-
cri¢ced and the total length was measured by means
of a calliper. The mean total body length was 1.98 cm
( 0.23, n 5 30) in the hatchery sample, and 1.93 cm
( 0.25, n 5 30) in the wild sample, and there was no
statistically signi¢cant di¡erence between the two
groups (t-test, 60, P40.1). Both hatchery and wild ju-
veniles spent a total period of 15 days in the holding
tanks before commencing the experimental trials.
Experimental procedures
The experimental tank was a 15.62 L (25 cm

25 cm
25 cm), completely bare, glass aquarium
¢lled with salt water (salinity 20) to a depth of
20 cm. Sheets of thick, dark-blue-coloured paper, si-
milar to those used in the housing tanks, covered
the rear and lateral walls of each aquarium to mini-
mize visual disturbance and provide an adequate vi-
sual contrast with the ¢sh silhouette to facilitate
video-tape analysis. A 25
25 grid of square cells
(1cm
1cm) was drawn on the external side of the
rear wall of the tank using a black marker (Fig. 1a).
This grid was used as a reference frame for beha-
vioural analysis. The experimental tank was
277
Antipredator reaction in sea bass juveniles S Malavasi et al.
(a)
d
c b
a and was operated using remote controls. Special care
e avoid disturbance of the experimental ¢sh caused
(b) b Experimental procedures for each experiment
d were conducted as follows:
c
a
e wild) were randomly selected from their housing
Figure 1 (a) Front view (not in scale) of the experimental
setup used in the ¢rst experiment (shadow). a, experimen-
tal tank; b,model bird predator; c, wire; d, lamps; e, electric
motor. (b) Lateral view (not in scale) of the experimental
setup used for the second experiment (dummy bill). a, ex-
perimental tank; b, dummy bill original position (not visi-
ble to the ¢sh); c, dummy bill stimulus position; d, lamp; e,
electric motor.
illuminated by suspending two 60 W lamps 40 cm
above the water surface.
Water temperature was measured before and at
the end of each experimental trial and averaged
20 1C, range  1 1C.
The ¢rst experiment (Fig.1a) was designed in order
to simulate a piscivorous bird £ying close to the water
surface. To achieve this, a wire was stretched over the
experimental aquarium, which allowed a rectangu-
lar table (51.5 cm length, 4.5 cm wide and 1cm thick)
to traverse 39 cm above the experimental tank just
below the lamps, creating a rapidly moving shadow
over the experimental aquarium. The model was
drawn by means of an electric motor with a velocity
of 85 cm s 1.
The second experiment (Fig. 1b) was designed
according to Giles (1984) and Wibe, Nordtug and
Jenssen (2001) in order to simulate a sudden over-
head attack from a piscivorous bird searching for
prey. We used a mechanically driven dummy bill
made of transparent solid PlexiglasTM (Venice, Italy)
that was lowered over the experimental tank, allow-
ing the tip of the bill to splash the water surface be-
fore raising it back again to its original position.
In both cases, the experimental apparatus used
to generate predator stimuli was not visible to the
278 Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 276^282
Aquaculture Research, 2008, 39, 276^282
experimental ¢sh before the moment of truth (MOT;
i.e. the point in time of presentation of the model, ac-
cording to Giles 1984 and Giles & Huntingford 1984)
was taken to isolate the electrical motors acoustically
and to position them in a separate surface in order to
both by noise and vibrations.
Experiment 1 ^ reaction to an overhead visual stimulus
(shadow)
Ten juveniles from each stock (i.e. hatchery-reared or
tanks, introduced into the experimental tank and
allowed to acclimatize for a 30-min period. Prelimin-
ary trials showed that after 30 min of acclimatiza-
tion, ¢sh resumed the ordinary swimming activity,
indicating that this time period was su⁄cient to al-
low recovery from the stress related to dip netting
and the transfer. At the end of the acclimatization
period, video recording of ¢sh behaviour commenced
and the model bird predator was drawn by the elec-
tric motor so as to pass above the experimental tank.
Recording of ¢sh behaviour continued for 1min after
the MOT. Overall, 10 groups of hatchery-reared and
wild juveniles were tested.
Experiment 2 ^ reaction to an overhead mechanical
stimulus (dummy bill)
Ten juveniles from each stock (i.e. hatchery-reared or
wild) were randomly selected from their housing
tanks, introduced into the experimental tank and
allowed to acclimatize for a 30-min period. At the
end of the acclimatization period, video recording of
¢sh behaviour commenced, and then the dummy bill
was lowered over the aquarium, allowing its tip to
splash the water surface before it was raised back to
its original position.Video recording of ¢sh behaviour
continued for an additional 1min (post stimulus per-
iod) after the MOT. Overall, four experimental groups
of hatchery-reared juveniles and four experimental
groups of wild juveniles were tested.
At the end of each experimental trial, all tested
individuals were transferred to empty housing tanks
and were not used for any other experimental trial.
Consecutively, the experimental tank was emptied,
rinsed and ¢lled with fresh salt water. All experimen-
tal trials were video recorded using the same digital
videocamera (Camcorder Canon MV400, Tokyo,
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Aquaculture Research, 2008, 39, 276^282
Japan), which was positioned in front of the experi-
mental tank before ¢sh transfer. Furthermore, we po-
sitioned a U-shaped opaque box behind the camera to
prevent visual contact between the experimental ¢sh
and the operator.
Behavioural variables and statistical analysis
All videotaped sequences were exported to a PC
and analysed using image analysis software (IMAGE
PRO-PLUS 3). Because this method analysed only the
two-dimensional aspect of visual images on video-
tapes, variations in the position of ¢sh along the third
dimension (i.e. from the front to the back of the tank)
were not compensated for.
The antipredator response of the shoal was
assessed using two independent measures: the dis-
tance from the tank bottom (DB10) and the freezing
duration (FD). The distance of the shoal from the tank
bottom was calculated to the nearest 0.01cm as the
average of the individual ¢sh distances from the
aquarium bottom at a given temporal frame for all
experimental trials (10 s after the MOT).
The duration of freezing was measured in ¢ve indi-
viduals (focal animal sampling; Altman 1974) ca-
sually selected from each replicate using image
analysis software (IMAGE PRO-PLUS 3). Freezing was
measured as a bout of behaviour starting when the
juvenile ceased all activities and rested immobile.
This technique made it possible to perform a ‘frame-
by-frame’ processing of the acquired videos providing
a 0.04 s accuracy. The operator, using the tagging
features provided by the software, was able to mea-
sure the individual freezing duration of ¢ve casually
selected individuals in each replicate. At the temporal
frame corresponding to the MOT, all ¢sh were
progressively numbered from the left to the right
and the freezing duration was measured in odd-
numbered ¢sh.
Table 1 Summary of results obtained in the two antipredator experiments
Variable Stimulus Hatchery
DB10 Shadow 1.29 (0.24–2.72)
DB10 Mechanical 1.91 (1.28–2.80)
FD Shadow 7.15 (4.70–9.78)
FD Mechanical 8.37 (6.16–11.77)
Mean values and range (in parentheses) of each variable for hatchery-reared and wild juveniles are reported together with statistical
signi¢cance of the comparison.
Signi¢cant at Po0.05.
DB10, distance from bottom measured 10 s after the Moment of Truth (cm); FD, freezing duration (s); NS, not signi¢cant.
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Antipredator reaction in sea bass juveniles S Malavasi et al.
Individual freezing duration and distance from
the bottom values were averaged for each replicate
and were tested for between-stock di¡erences
using the Mann^Whitney U-test. All statistics
were carried out by means of STATISTICA 6.0 software
package.
Results
Videotape analysis con¢rmed that sea bass juveniles
engaged an escape (£eeing) and freezing reaction
once presented with the stimulus. In some cases,
animals from both groups engaged an escape reac-
tion (£eeing) towards the bottom (see below and Table
1 for mean values) and then a freezing one, while in
other cases individuals (mostly those close to the bot-
tom at the MOT) commenced a freezing reaction as
soon as presented with the stimulus. After a variable
period spent motionless at the bottom, individuals
resumed the ordinary swimming activities observed
before the MOT.
The mean distance from the bottom (10 s after the
MOT) varied from ca. 1.3 to 1.8 cm in the ¢rst experi-
ment (shadow; Table 1) for shoals composed of hatch-
ery-reared and wild juveniles respectively. For the
second experiment (mechanical stimulus; Table 1), the
mean distance from the bottom was ca. 1.9 and 1.7 cm
for the shoals composed of hatchery-reared and wild
juveniles respectively. The mean values were therefore
similar for both groups in both experiments and there
was no statistically signi¢cant di¡erence (Table 1).
The mean freezing duration varied from ca.7 s in the
hatchery-reared shoals to ca. 10.5 s in the wild shoals
as regards the ¢rst experiment (shadow; Table 1,
Fig. 2a). For the second experiment (mechanical
stimulus, Table 1), the mean freezing duration was ca.
8.5 s and ca. 13.5 s for the shoals composed of hatch-
ery-reared and wild juveniles respectively (Fig. 2b).
Wild n Mann^Whitney U-test
1.83 (0.35–4.44) 10 NS
1.69 (0.62–3.59) 4 NS
10.40 (5.97–13.00) 10 
13.74 (10.40–16.57) 4 
279
Antipredator reaction in sea bass juveniles S Malavasi et al. Aquaculture Research, 2008, 39, 276^282

(a) 16 the freezing duration tended to be longer in wild ¢sh

14
than in sea-ranched ¢sh. Furthermore, a similar
quantitative di¡erence in freezing behaviour, as a
Freezing duration (sec)

12 response towards aerial predation, was found by

10 Templeton and Shriner (2004) between two guppy
populations, characterized by di¡erent habitat struc-
8
tures and predation regimes. These two cases, to-
6 gether with the results of the present study, suggest
that freezing is a fundamental component of the anti-
4
predator behavioural response of ¢sh towards aerial
2 stimuli and that the duration of this behaviour can
0 show a high degree of intra-speci¢c variation. It has
Hatchery reared Wild been shown that freezing is a clear and discriminating
(b) 16 behaviour in the antipredator tests, as it proved to be
14 responding highly towards several chemical and vi-
sual stimuli (Giles 1984; Giles & Huntingford 1984;
Freezing duration (sec)

12
10
8
6 Vilhunen & Hirvonen 2003). As shown by our results
4 and those obtained by Petersson and Jarvi (2006),
2
0 ing of £eeing towards the bottom and freezing (mo-
Hatchery reared Wild
Figure 2 Means and standard errors of the freezing
duration (sec) for each group (hatchery vs. wild). (a) Over-
head visual stimulus (shadow, 10 replicates). (b) Overhead
mechanical stimulus (dummy bill, four replicates).
In both experiments, the mean freezing duration
was signi¢cantly higher for the experimental groups
composed of wild juveniles (Table 1), than that com-
posed of hatchery ¢sh.
In conclusion, the results showed that shoals
of wild juveniles di¡ered from those formed by
hatchery-reared ¢sh in the mean freezing duration
induced by the presentation of the stimulus.
Discussion
This study showed that hatchery-reared and wild sea
bass juveniles reacted similarly towards overhead
fright stimuli (both visual and mechanical) but their
responses di¡ered quantitatively, with wild juveniles
freezing signi¢cantly longer than hatchery ones.
These results are consistent with those found by
Petersson and Jarvi (2006) in sea-ranched, wild and
crossed brown trout. These authors found that wild
¢sh responded stronger to a heron dummy and that
Giacquinto & Volpato 2001; Vehanen 2003; Vilhunen
& Hirvonen 2003; Lehtiniemi 2005). It is therefore
likely that this response has strong innate determi-
nants, as suggested by some authors (Giles 1984;
the same antipredator behavioural sequence was
exhibited by both wild and hatchery juveniles, consist-
tionless period), followed by the recovery of the
ordinary activities. This suggests that this behavioural
sequence is basically innate and inherited, as also hy-
pothesized by Giles (1984) for laboratory-reared
sticklebacks.
Nevertheless, the quantitative di¡erences evi-
denced by both the present and the previous results
indicate that reared ¢sh show less intense responses
towards overhead fright stimuli than wild ¢sh, sug-
gesting that experience could play a role in generat-
ing this pattern. This also agrees with the results of
Johnsson, Hojesjo and Fleming (2001), Johnsson,
Sernland and Blixt (2001), who found that wild
young salmon showed a more intense escape reac-
tion than hatchery ¢sh of the same genetic origin to-
wards a dummy heron bill. As pointed out by several
authors, the lack of some key experiences in the
hatchery environment may generate behavioural dif-
ferences at every life stage (Price 1999; Kelley &
Magurran 2003; Huntingford 2004). In particular,
the role of habituation has also been often outlined
in relation to the responses of ¢sh to farm workers
(Kelley & Magurran 2003; Huntingford 2004). Farm
workers often constitute the only overhead fright sti-
mulus to which hatchery ¢sh are subjected and it is
likely that ¢sh become rapidly habituated to that,
as this stimulus is not e¡ective in the hatchery.

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280 Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 276^282
Aquaculture Research, 2008, 39, 276^282
Habituation could therefore be one of the possible ex-
periential factors generating the di¡erence we found
between hatchery and wild sea bass juveniles. There
are no data available in the current literature showing
whether the e¡ects of repeated disturbance and con-
sequent habituation due to the farm operators in the
hatcheries based on mesocosm technique are weaker
than that in the hatcheries using more traditional,
intensive systems. In our investigated system, farm
workers were used to operate repeatedly on the border
of the large rearing tanks for routine management
and for this reason it is possible to hypothesize that
habituation was acting on the hatchery juvenile ¢sh.
In a previous study by Malavasi et al. (2004), di¡er-
ences between wild and hatchery sea bass juveniles
were shown in terms of behavioural responses to-
wards a live ¢sh predator (a common eel). Also, in
that case, di¡erences were related to some quantita-
tive features of the antipredator responses, the beha-
viours of the two juvenile groups qualitatively being
very similar. It therefore appears that the behavioural
responses to aquatic predation parallel those exhib-
ited towards aerial predation as shown also by Tem-
pleton and Shriner (2004) for the guppy populations.
In both the present work and that by Malavasi et al.
(2004), wild and hatchery juveniles originated from
two di¡erent populations, so that genetic di¡erences
could not be ruled out completely. However, for both
aquatic and aerial predation, there is no evidence of
di¡erential predation pressures between the Venice
lagoon and the Southern coasts of Italy (where wild
juveniles were caught). Furthermore, it is likely that
bird predation is widespread and common in every
coastal and lagoon environment, suggesting that
the observed di¡erences between the two groups of
juveniles could be due to the e¡ects of early environ-
ment rather than genetic factors related to popula-
tion origin. According to Salvanes and Braithwaite
(2006), behavioural de¢ciencies generated in early
life are likely to a¡ect behaviour in later stages. Our
results suggest that the experience-deprived hatch-
ery environment may have generated the less intense
behavioural response of hatchery-reared sea bass ju-
veniles, especially through habituation mechanisms,
and this could have negative e¡ects on the later
success of the ¢sh. As shown by Salvanes and
Braithwaite (2006), environmental enrichment of
rearing tanks could generate more £exible beha-
viours in hatchery cod, potentially improving their
skills later. It seems that behavioural de¢cits gener-
ated by learning processes related to the hatchery en-
vironment are a common pattern for marine and
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Journal Compilation r 2008 Blackwell Publishing Ltd, Aquaculture Research, 39, 276^282
Antipredator reaction in sea bass juveniles S Malavasi et al.
freshwater-cultured ¢sh, and it is therefore very use-
ful to further investigate the e¡ects of environmental
enrichment on their behaviour.
Acknowledgments
We are grateful to Professor Stefano Cataudella and
Dr Attilio SpanoØ for having provided the hatchery
and experimental facilities for the present study. We
would also like to thank Mr Piero Trebbi for the tech-
nical support provided for the experimental set-up.
This study was funded by the Italian Ministry of
Agricultural Resources (MIPAF, VI Piano triennale
della pesca e dell’acquacoltura:‘Ripopolamento attivo
di lagune, stagni costieri, e localizzate aree della fas-
cia costiera con giovanili certi¢cati di specie ittiche
e di crostacei, secondo i principi del codice di condot-
ta per una pesca responsabile (FAO 1995)’).
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