Aquaculture Research, 2011, 42, 441^449 doi:10.1111/j.1365-2109.2010.02639.

The effects of tank colours on the growth and survival

of cuttlefish (Sepia officinalis, Linnaeus 1758)
hatchlings and juveniles

Antonio V Sykes1,2, Pedro M Domingues3, Lorenzo Marquez2 & Jose P Andrade1

1
CCMAR-CIMAR L.A., Centro de Ciencias do Mar do Algarve, Campus de Gambelas, Universidade do Algarve, Faro, Portugal
2
IFAPAAgua del Pino (Junta de Andaluc| a), Huelva, Spain
3
IEO ^ Centro Oceanograco de Vigo, Cabo Estai, Canido,Vigo, Spain

Correspondence: AV Sykes, CCMAR-CIMAR L.A., Centro de Ciencias do Mar do Algarve, Campus de Gambelas, Universidade do Algarve,
8005-139, Faro, Portugal. E-mail: asykes@ualg.pt

Abstract
The eects of dierent tank colours (white, yellow
sandy and black) on the growth, mortality and
biomass production were studied for hatchling and
early juvenile cuttlesh. For hatchlings, the use of
dierent colour tanks did not promote dierences in
growth due to the higher variability (standard devia-
tion) found in the white- and sand-coloured tanks.
Black tanks promoted the lowest and highest values
for total mortality and biomass respectively. For
juveniles, the use of dierent tank colours promoted
dierent growth (Po0.05), but not mortality. Black
tanks promoted the best results in terms of growth
and biomass. The results obtained in the present
study advise the use of black (or dark colour) tanks
in the hatchling and early juvenile stages to reduce
the standard deviations associated with growth,
mortality and biomass production. This will contri-
bute to minimize problems associated with slow and
fast growers and competition.
Keywords: biomass production, culture stage,
European cuttlesh, Sepia ocinalis, tank colours
Introduction
The cuttlesh (Sepia ocinalis, Linnaeus) is one of the
most well-known cephalopods (Boletzky 1983) and a
good candidate to become one of the rst cephalopod
species in commercial aquaculture (Boucaud-Camou
1990; Barnabe 1996). Recently, Sykes, Domingues,
Correia and Andrade (2006) reviewed the aquacul-
ture potential, the state of the art and future trends
of research for the species. In this work, the authors
highlighted a series of bottlenecks that need to be re-
solved before the technology can acquire a maturity
that will enable a transition to an industrial scale,
one of it being the problem related to survival in the
hatchling stage. It is known that the rst-stage cul-
ture of this species is the most problematic, due to de-
pendence on live food (Sykes, Domingues, Correia
et al. 2006). Feeding is the most crucial factor deter-
mining both survival and development, and its im-
portance is enhanced by temperature (Domingues,
Bettencourt & Guerra 2006; Sykes, Domingues & An-
drade 2006) and competition. Nonetheless, other re-
lated eco-biological factors should also be of great
importance to achieve good results. In fact, system
design may aect the success of culturing cuttlesh
(Lee, Turk, Forsythe & DiMarco 1998), promoting
stress and lower animal welfare as described by
Conte (2004) for nsh.
In order to enhance growth and survival rates dur-
ing the post-hatching phase of S. ocinalis, a number
of experiments related to the surrounding environ-
ment under culture conditions have already been
conducted by Boletzky (1996), Dickel, Boal and Budel-
mann (2000), Sykes, Domingues, Loyd, Sommereld
and Andrade (2003) and Poirier, Chichery and Dickel
(2004). Lee et al. (1998) stated that tank colour is
extremely important in determining hatchling survi-
val in squid, while it is not so important in sepia. In
fact, the use of black walls seems to be benecial in
terms of stress and well-being to other cephalopod
species, because it increases contrast and prevents

r 2010 Blackwell Publishing Ltd 441

Optimizing cuttlesh growth and survival through tank colours AV Sykes et al. Aquaculture Research, 2011, 42, 441^449
wall collision in squid (Hanlon, Hixon & Hulet 1983;
Vidal, Dimarco,Wormuth & Lee 2002). Nabhitabhata
and Nilaphat (2000) were pioneers in applying the
concept of tank colour as a growth and survival
enhancer for cephalopods, in juvenile Sepia pharaonis
culture. In their work, in an attempt to determine the
preference for brightness, they used white, grey and
black tanks. Sepia pharaonis individuals showed a
preference for low levels of brightness, which were
associated with rst grey and then black tanks.
Nevertheless, the same authors also stated that
this preference was highly species dependent, and
on particular physiological needs, and the study was
not growth related and rather choice driven.
In sh species, tank colour is used to enhance
growth and survival, and diminish mean weight
standard deviations under culture conditions. For in-
stance, the reectance and colour of tank walls was
shown to increase the prey capture rates in larval
haddock (Downing & Litvak 1999) and grouper (Dur-
ay, Estudillo & Alpasan 1996), thus inuencing both
growth and survival. Nonetheless, until now, the in-
uence of tank colours had not been studied in the
European cuttlesh, being animals usually reared in
white tanks in our culture facilities.
Therefore, the main question is whether tank col-
our would somehow inuence cuttlesh growth, the
mean weight standard deviations and mortality,
and whether this colour should be black to resemble
darkness (night conditions) or whether it should be
sandy-like to resemble the natural surrounding con-
ditions in the wild during daylight. Consequently, the
objective of the present study was to determine how
dierent colours (white, sand or black for hatchlings,
and black or white for juvenile) aected the growth,
survival, tank biomass production of cuttlesh dur-
Table 1 Experimental conditions for both trials
Tank colour Black
Experiment I
Temperature (all)
Salinity (all)
Dissolved oxygen (all)
Incident light intensity (all)
Reflective light intensity 10.63  1.27 lxa
Experiment II
Temperature (all)
Salinity (all)
Dissolved oxygen (all)
Incident light intensity (all)
Reflective light intensity 3.80  0.62 lxa
Superscript letters represent the grouping of dierences within the same row for Po0.001.
442
ing the rst 40 DAH (fed live prey), and juveniles from
30 to 80 DAH (with a transition of live to frozen prey
as part of the commonly used weaning protocol).
Material and methods
Experiment I ^ hatchlings
The experiment lasted for 40 days, corresponding to
the hatchling stage. A total of 180 cuttlesh hatchl-
ings with a mean wet weight (MWW) of 0.084 
0.015 g were used. All cuttlesh hatched in the same
day, from a single brood obtained from cultured
females reproducing in the facilities of the IFAPA
Agua del Pino Experimental Aquaculture Station
(Cartaya, Huelva, Spain). The embryonic develop-
ment of eggs as well as broodstock rearing followed
the protocols described in Sykes, Domingues and An-
drade (2006).
A ow-through system composed of nine similar
rectangular tanks was used. Each tank had a total vo-
lume of 10 L and a bottom area of 0.141m2. The water
ow was set to 17 L h  1. Water was pre-ltered
through an industrial sand lter, decanted and, be-
fore entering the tanks, passed through a 25 mm mesh
lter and a UV bacteriological lter. Water tempera-
ture, salinity and dissolved oxygen were measured
daily in the morning. Both temperature and dissolved
oxygen were measured using a CRISON OXI45 (Bar-
celona, Spain) probe, while salinity was measured
using a Kikuchi optical salinity meter. The mean cul-
ture temperature, salinity and dissolved oxygen con-
ditions are presented in Table 1. A natural daylight
cycle was provided (14 h L:10 h D), which corre-
sponded to the local natural conditions in spring. In-
cident and reective light intensity were measured
Sand White
19.3  0.5 1C
36.7  0.9 g L  1
7.4  0.1 mg/L (98.2  1.3%)
320.00  30.00 lx
124.03  11.89 lxb 130.77  4.18 lxb
20.5  1.0 1C
38.2  1.0 g L  1
7.0  0.4 mg/L (95.0  5.2%)
320.00  30.00 lx
46.53  9.00 lxb
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449
Aquaculture Research, 2011, 42, 441^449 Optimizing cuttlesh growth and survival through tank colours A V Sykes et al.
daily, using a Digital Light Meter ITALIA HT-170 (HT
Italia, Faenza, Italy), and the mean recorded values are
presented in Table1. Dierences in light reection were
only found between the black tanks and the remaining
two colours (Po0.001). To ensure these light condi-
tions, tanks were cleaned daily by suction with a small
air hose and with a sponge on sampling days. The
choice of the colours tested was made according to
the following: white ^ commonly used in the eld sta-
tion of the Ramalhete (CCMAR, University of the Al-
garve, Faro, Portugal) for cuttlesh culture in the past
10 years; yellow sandy ^ used to resemble a sandy bot-
tom without all its disadvantages (Sykes et al. 2003);
and black ^ in order to stimulate prey catching as sug-
gested by previous authors (Hanlon & Messenger1988;
Budelmann, Riese & Bleckmann 1991; Boletzky 1996).
All paint used in the present experiment was polyur-
ethane plus acrylic enamel based. Ink references were
from the ARALCIN paint line (CIN inks) and as follows:
black ^ C1 80-301.0531/06; sand ^ C1 80-301.0507/06;
and white ^ C180-301.0501/06.
Newly born cuttlesh hatchlings were used for the
three dierent-coloured tanks tested. Each replicate/
tank contained 20 hatchlings, thus not representing
a high density nor in the limit of bottom areas estab-
lished by Sykes et al. (2003), which generate stress
and mortality.
Because no grass shrimp was available near the
culture facilities, live mysids (Mesopodopsis slabberi)
and adult Artemia sp. were collected from ponds and
salt ponds respectively. Mysids were used as the
primary feeding prey, while Artemia sp. was only
added when there were not enough mysids to cover
the feeding necessities of all the individuals/tanks.
Cuttlesh were always fed ad libitum and, when
needed, with the same mysid^artemia proportions.
Experiment II ^ transition from hatchling to
the juvenile stage
A total of 210 cuttlesh juveniles, 30 days of age, and
with an MWWof 0.488  0.146 g were used. All cut-
tlesh used were born on the same day and from egg
clusters collected in the intertidal zone of the beach
of El Rompido (Cartaya, Huelva, Spain). Technology
and location similar to that described for experiment
I was used. After hatching and during the rst 30
DAH, cuttlesh were fed live mysids (M. slabberi) and
kept in similar white rectangular tanks.
Because no signicant dierences among tank col-
ours have been calculated for the previous experi-
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449
ment, only the colours producing the higher
dierence in light reection were used: white and
black. Also, because white and yellow sandy dis-
played similar light reection and the rst is com-
monly used in the rearing of this life stage, white
was chosen for this second trial.
A ow-through system composed of six similar
rectangular raceway-type tanks (three white and
three black) was used. This system was similar to that
of experiment I, except for the size, shape and depth
of the tanks. Each tank had a total volume of approxi-
mately 40 L and a bottom area of 0.284 m2. To prevent
cuttlesh from jumping out, all tanks were covered
with a 1mm plastic white mesh.
Water temperature, salinity, dissolved oxygen and
light intensities were monitored using the same pro-
cedure as described for experiment I and the re-
corded values are presented in Table 1. A natural
daylight cycle and light intensity at the air^water
tank interface similar to that of experiment I was
provided. Dierences in light reection were found
between black and white tanks (Po0.001).
Each replicate/tank contained 35 cuttlesh juve-
niles, thus not representing a high density nor in the
limit of bottom areas (Sykes et al. 2003).
During the rst 20 days of experiment, live mysid
shrimp (M. slabberi) was supplied ad libitum. After
that, frozen adult grass shrimp (Palaemonetes varians)
was supplied as food in a quantied ad libitum. There-
fore, cuttlesh were fed in excess but the amount sup-
plied was quantied and based on the remains that
were removed approximately 24 h later, i.e., the next
morning. The quantication was performed by the
overall cuttlesh weight (total biomass) present in
each tank, from growth data collected every 10 days.
Data collection and analysis
Before the start of each experiment, cuttlesh in each
tank colour were weighed and an ANOVA test (Zar
1999) was performed to ensure that no signicant
dierences were found among replicates. In both ex-
periments, all cuttlesh were weighed every 10 days.
Data collected were used to calculate (1) the MWW;
(2) the mean absolute instantaneous growth rate
(MAIGR) (%BWday  1) 5 (LnW2  LnW1)/t  100,
where W2 and W1 are the nal and the initial mean
weight for each colour treatment, respectively, Ln
the natural logarithm and t the number of days of
the total experiment; (3) the total absolute mortality
(TAM), which expresses all dead animals for each col-
443
Optimizing cuttlesh growth and survival through tank colours AV Sykes et al. Aquaculture Research, 2011, 42, 441^449

our treatment; (4) the mean cumulative mortality (%) Table 2 Mean absolute IGR (MAIGR), mean cumulative
(MCM) 5 (Sdead/initial live animals)  100 for each mortality (MCM), total absolute mortality (TAM), biomass
colour treatment; (5) biomass (B) (g) 5 S of WW of (B) and mean biomass increase (MB%) values for cuttlesh
all cuttlesh for each colour; and (6) the mean bio- hatchlings and juveniles at the end of both experiments
mass relative increase index (B%) (%BWday  1) 5
Tank colour Black Sand White
[(Bf  Bi)/(Bi  t)]  100, where Bf and Bi represent
the nal and the initial biomass for each colour treat- Experiment I
ment, respectively, and t is the number of days of the MAIGR 4.62  0.161 4.16  0.684 4.34  0.572
(%BW day  1)
total experiment.
MCM (%) 21.7  5.8 30.0  13.2 36.7  7.6
Results are presented as means  standard devia-
TAM (cuttlefish) 13 18 23
tion (SD) for triplicate tanks. At the beginning and the B (g) 25.04 18.40 17.63
end of the experiments, data were checked for a normal B% 9.74  0.93 6.78  2.57 6.22  1.74
distribution using the one-sample Kolmogorov^ (%BW day  1)
Experiment II
Smirno test (Zar 1999), as well as for homogeneity of
MAIGR 3.66  0.08 3.13  0.21
variances using Levenes test (Zar 1999). When neces-
(%BW day  1)
sary, logarithm or arcsin transformation of the data TAM (cuttlefish) 0 2
was performed. When a normal distribution and/or B (g) 318.3 242.2
homogeneity of the variances were not achieved after B% 10.48  0.52 7.43  1.26
(%BW day  1)
transformation, robust tests were used. Statistical dif-
ference was considered for Po0.05.
A nested ANOVA test (Zar 1999), withtank colouras resumes the statistical analysis of tank colour eect
a xed factor and tankas a random factor, was per- on these growth parameters. At the end of the experi-
formed to test for dierences in individual wet weight ment, all reared cuttlesh attained similar weights
on the rst and the last day of each experiment. Dif- (P40.05; Table 3) and all three tank colour groups
ferences were not found on the rst day (data not displayed exponential growth (Fig. 1).
shown). When dierences were found, a post hoc un- MAIGR was not statistically dierent among col-
equal N HSD test was used to determine the homoge- ours (P40.05;Table 3). Nevertheless, MAIGR in black
nous subsets. As for MAIGR, dierences were tanks displayed smaller SD when compared with
assessed through using aWelch test (Zar 1999) for ex- sand and white tanks (Table 2).
periment I, while a Students t test (Zar 1999) was Mortality among the dierent-coloured tanks, at
used for experiment II. the end of the experiment, was not statistically dier-
MCM dierences between colours were assessed ent in terms of MCM (Fig. 2; P40.05; Table 3), but
using one-way ANOVA and Students t test (Zar 1999), TAM displayed dierent values among colours, being
in experiments I and II respectively. lower in the black treatment (Table 2).
Dierences in B and B% were assessed in both ex- Regarding B, cuttlesh reared in black tanks at-
periments using a repeated measures one-way ANOVA tained the higher biomass,1.42 times higher than that
(Zar1999).The Greenhouse^Geisser correction was ap- obtained for those reared in the white tanks (Table 2).
plied to the calculation of the degrees of freedom when Black tanks displayed a mean B of 8.35  0.99 g, while
the tests did not t the sphericity assumption. When yellow-sandy tanks showed 6.13  1.75 g and white
dierences were found, a post hoc Tukey test (Zar tanks attained 5.88  1.28 g. B% did not display dier-
1999) was used to determine the homogenous subsets. ences (P40.05) between tank colours at the end of the
To adjust the signicance level, while performing experiment (Table 3). However, B% was always higher
several hypothesis contrasts, the stepwise Bonferroni in black tanks than the tanks of the other colours, but
and the step-up FDR corrections (Zar 1999) were ap- not statistically dierent (P40.05) due to the higher va-
plied (Table 3), with similar results. lues of SD found in both sand and white tanks (Table 2).

Results
Experiment I ^ hatchlings
Table 2 shows the results obtained in terms of the va-
lues of MAIGR, MCM,TAM, B and B%, while Table 3
Experiment II ^ transition from hatchling to
the juvenile stage
Table 2 resumes the results obtained in terms of
MAIGR, TAM, B and B%, while Table 3 resumes the

444 r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449

Aquaculture Research, 2011, 42, 441^449 Optimizing cuttlesh growth and survival through tank colours A V Sykes et al.

Table 3 Statistical analysis of the eect of tank colour on dierent growth parameters of Sepia ocinalis in experiments1and 2

Homogeneous
Experiment Variable Test Source SS d.f. Statistic P Post hoc groups

Experiment 1 WWDay 40 Nested Colour 0.206 2.00 5.07 0.145

ANOVA Tank (Col.) 0.230 6.00 1.89 0.088
B RM ANOVA Colour 7.990 2.00 2.73 0.144
Time 168.0 1.06 133.60 o0.001 Tukey D1, D10, D20o
D30oD40
Interact. 7.500 2.12 2.99 0.121
MAIGR Welch test Colour NS 2.00 3.52 0.173
B% RM ANOVA Colour 0.029 2.00 2.72 0.144
Time 0.770 1.42 48.11 o0.001 Tukey D10oD20o
D30, D40
Interact. 0.180 2.85 0.56 0.650
p
Arcsin MCM ANOVA Colour 0.042 2.00 1.81 0.242
Experiment 2 Log (WWDay 50) Nested Colour 0.592 1 30.54 0.005 Unequal-N WhiteoBlack
ANOVA HSD
Tank (Col.) 0.077 4 0.76 0.552
B RM ANOVA Colour 849 1 41.60 0.003 Tukey WhiteoBlack
Time 24827 5 482.00 o0.001 Tukey D1oD10oD20o
D30oD40oD50
Interact. 720 5 14.00 o0.001
MAIGR RM ANOVA Colour 2.18 1 17.96 0.013 Tukey WhiteoBlack
Time 41.76 2.23 60.09 o0.001 Tukey D204D104D30,
D 40, D50
Interact. 0.369 2.23 5.53 0.624
B% RM ANOVA Colour 5.078 1 14.85 0.018 Tukey WhiteoBlack
Time 58.68 2.70 58.18 o0.001 Tukey D10, D204D30,
D50  D40, D50
Interact. 1.134 2.70 1.12 0.377
p
Arcsin MCM t-test Colour 4  1.00 0.374

d.f. was calculated after applying the Greenhouse^Geisser correction for sphericity.
Black bold P-values are signicant after a stepwise Bonferroni or a step-up FDR correction; grey bold P-values are signicant after a
step-up FDR correction.
NS, not supplied by the SPSS software; RM ANOVA, repeated measures ANOVA; Tank (Col.), the variable Tank nested into the variable
Colour.

statistical analysis of tank colour eect on these

Figure 1 Growth in the mean wet weight (g) of hatchl-
ings of dierent-coloured groups (B, black; S, sand and W,
white) and exponential growth curves with respective
equations for experiment I (Exp. B, black; Exp. S, dark grey;
and Exp.W, light grey). Data are presented in oset.Vertical
lines represent the standard deviation.
growth parameters. After the rst 20 days and until
the end of the experiment, cuttlesh, cultured
in both groups and fed ad libitum, were found to be
consuming the same amount of food, which was ap-
proximately 25% of their body weight per day. In this
life stage, the use of dierent tank colours promoted
dierent growth (Po0.05) between colour treatments
(Table 3) and growth was linear (Fig. 3). At the end of
the experiment, cuttlesh reared in black tanks
weighed more (Po0.05) than those of white tanks
(Table 3). MAIGR was also dierent (Po0.05) between
colours at the end of the experiment (Table 3).
Mortality was only observed in a white tank (TAM:
two individuals ^ 6% for the tank; Table 2) and was
not statistically dierent in terms of MCM (P40.05;
Table 3).

r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449 445

Optimizing cuttlesh growth and survival through tank colours AV Sykes et al. Aquaculture Research, 2011, 42, 441^449
Figure 2 Mean cumulative mortality [MCM (%)] of
hatchlings of dierent-coloured groups (B, black; S, sand
and W, white) for experiment I. Data are presented in o-
set.Vertical lines represent the standard deviation.
Figure 3 Growth in mean wet weight (g) of juveniles of
dierent-coloured groups (B, black and W, white) and lin-
ear growth curves with respective equations for experi-
ment II (Exp. B, black and Exp. W, dark grey). Data are
presented in oset. Vertical lines represent the standard
deviation.
Dierences were found in B (Po0.05; Table 3),
where black tanks attained the higher biomass (Table
2). Black tanks displayed a mean B of 106.1  5.0 g,
while white tanks attained 80.7  7.5 g. Accordingly,
B% displayed dierences (Po0.05) between colours
at the end of the experiment (Table 3), and was high-
er in black tanks (Table 2).
Discussion
A methodological point: biomass production
as a new variable
This study intended to incorporate two culture stages
(hatchling and juvenile) and their transition (which
446 r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449
is food dependent), in view of the fact that the species
usually display dierent growth patterns for each
one. The main eects that allow a correct colour eva-
luation are growth and mortality, because they will
ultimately aect biomass production. The present
study also introduces two new concepts of measur-
ing the growth of cephalopods while conducting stu-
dies in captivity, which are B and B%.
According to Forsythe and Van Heukelem (1987)
and Forsythe (1993), the cephalopod hatchling stage
is characterized by exponential growth, while the ju-
venile stage commonly displays logarithmic growth.
MAIGR is commonly used in cephalopod growth ex-
periments, which is based on mean weights and
could overlook the eect of mortality. Therefore, we
have introduced an additional index, B%, based on
the total biomass present in the tanks. This is particu-
larly important because companies that produce a
particular species are selling a given biomass (B)
and not a mean weight. Another key aspect is that
reared animals should have a similar weight that
(with a lower standard deviation between animals)
derives from lower competition for food, among other
reasons. The new tools (B and B%) introduced in the
present study are seen as useful to provide a more ac-
curate vision and prediction over tank production of
the species.
Eects of colour choice on the growth of
hatchlings and juvenile cuttlesh
We believe that the lack of dierences found in experi-
ment I was mainly due to the high variability (higher
standard deviation ^ Table 2) found in both white and
sand tanks. In fact, the use of black tanks in this stage
led to less variable values of mortality and biomass
gain. As for experiment II, dierences were found be-
tween colours for all the indexes used, except for
mortality.
Most of the mortality in cuttlesh culture occurs at
the hatchling stage (Sykes, Domingues, Correia et al.
2006) and this may be caused by several factors al-
ready identied, such as the amount and quality of
prey (Domingues, Kingston, Sykes & Andrade 2001;
Domingues, Sykes & Andrade 2001; Domingues,
Sykes, Sommereld, Almansa, Lorenzo & Andrade
2004), or minimum bottom areas (Sykes et al. 2003),
and others. We intended to understand the eects of
tank colour on cuttlesh growth and survival but,
while analysing the results, we realized that those
cannot be dissociated from light intensity (which
Aquaculture Research, 2011, 42, 441^449 Optimizing cuttlesh growth and survival through tank colours A V Sykes et al.
was similar among tanks for incident light but com-
pletely dierent for reective light). In fact, black
tanks displayed a signicantly lower reective light
intensity than the remaining tested colours, and the
present results reect this dierence. The present re-
sults were dierent from those obtained by Nabhitab-
hata and Nilaphat (2000) for S. pharaonis in terms of
colour. Nonetheless, grey was not tested in the pre-
sent study and neither was tank colour evaluated in
that study. We believe that a combination of black
tanks and incident light intensity might promote
adequate brightness and contrast conditions and
stress the importance of conducting experiments re-
garding brightness and contrast by using dierent
light intensities. This is due to the fact that, according
to the latter authors, brightness and contrast will in-
terfere with visual discrimination and, therefore,
hunting.
Theoretical relationship between cuttlesh
visual acuity and predation behaviour
While developing culture systems for a given species,
its biology and ecology must be considered. The Eur-
opean cuttlesh is a visual hunter (Messenger 1968)
and has an excellently developed neural anatomy,
which is responsible for its distinctive sensory abil-
ities, which include mechanoreception, chemorecep-
tion and vertebrate-like vision (Budelmann & Young
1993; Boal, Shashar, Grable,Vaughan, Loew & Hanlon
2004), all of them contributing to prey hunting.
Cuttlesh eyes display high sensitivity to light and
a visual acuity that is far better than that of sh
(Watanuki, Kawamura, Kaneuchi & Inashita 2000),
which is due to their photosensitive pigments (Wells
1966; Hara 1968). The degree of cuttlesh pupil con-
striction is ambient light intensity dependent (Dou-
glas, Williamson & Wagner 2005) and, during the
present study, reective light in both sand and white
tanks was10 times higher than that in black tanks. In
fact, the retinal structures of cuttlesh are ideal for
optimal light capture in dark environments (Davies,
Gowen, Krebs, Schertler & Saibil 2001) and do not re-
spond well to bright light, as they lack a consensual
light-induced pupil response (Douglas et al. 2005).
The fact that the species is colour blind (Marshall &
Messenger 1996; Mthger, Barbosa, Miner & Hanlon
2006) does not interfere with the quality of its vision,
because adequate visual information is gathered by
contrasting light and dark shades (Boal et al. 2004).
According to Mthger et al. (2006), cuttlesh is able
r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449
to perceive objects in their background that dier in
contrast by 15%. At the time of feeding, while indivi-
duals in black tanks remained quiet and camouaged
in the bottom, waiting for the prey to get into their
visual striking area, white and sand individuals
would move more in the tanks looking for the prey.
According to Marshall and Messenger (1996), the
species neural anatomy allows it to camouage with
the surrounding environment, because it is able to
perceive dierent light intensities in the background
and generate contrast-dependent patterns. This abil-
ity is used by cuttlesh for hunting and concealment
(camouage) and has been described extensively by
other authors, such as Hanlon and Messenger (1988,
1996).
In experiment I, when the size dierences among
the individuals increased, larger cuttlesh present in
both white and sand tanks were repeatedly seen to
compete with smaller cuttlesh after these had
caught a prey, leading the smaller individuals to re-
lease this captured food. This competition has prob-
ably led to even higher dierences in size that
eventually resulted in starvation and mortality.
Nevertheless, in experiment II, because dead prey
remained quiet at the bottom of the tank, cuttlesh
present in the black tanks theoretically had to spend
the same amount of energy as those in white tanks
for hunting. Therefore, the dierences found may be
related to the capacity of cuttlesh to see the prey, i.e.
the brightness and contrast of the tanks, which is
light intensity related.
The use of dierent tank colours, with a similar in-
cident light, produced distinct reective light intensi-
ties in the tanks, thus producing dierent brightness
and contrast in the tanks.
Final considerations
Black tanks facilitated easier tank cleaning because
the remains of dead and uneaten prey were more visi-
ble than in the other tanks, especially while using
grass shrimp. Generally speaking, this colour im-
proved growth and survival, which were translated
into higher biomass increase and more accurate and
precise growth values. Also, the use of black tanks in
the hatchling and juvenile stages can be used as a so-
lution to reduce the problem with slow and fast
growers resulting from competition. Nevertheless,
there is a need for further studies regarding the use
of black tanks with other culture variables such as
at the adult reproducing stage, when an articial diet
447
Optimizing cuttlesh growth and survival through tank colours AV Sykes et al. Aquaculture Research, 2011, 42, 441^449
is developed, and culture at high densities in every
stage of the life cycle. Additional studies are also ne-
cessary to clarify the inuence of incident and reec-
tive light intensity in every culture stage and using
black tanks.
Acknowledgments
Antonio Sykes wishes to thank the Fundacao para a
Ciencia e a Tecnologia, which funded the current re-
search by way of a PhD (SFRH/BD/12409/2003) and
a Post-Doc (SFRH/BPD/36100/2007) grant. The rst
author is also indebted to the IFAPA, from the Junta
de Andalucia, which allowed him to conduct the ex-
periments at the CIFAP Agua del Pino Aquaculture
Research Facilities.We also wish to thank two anon-
ymous referees for their valuable criticism.
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