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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 ture potential, the state of the art and future trends
of research for the species. In this work, the authors
The eects of dierent tank colours (white, yellow
highlighted a series of bottlenecks that need to be re-
sandy and black) on the growth, mortality and
solved before the technology can acquire a maturity
biomass production were studied for hatchling and
that will enable a transition to an industrial scale,
early juvenile cuttlesh. For hatchlings, the use of
one of it being the problem related to survival in the
dierent colour tanks did not promote dierences in
hatchling stage. It is known that the rst-stage cul-
growth due to the higher variability (standard devia-
ture of this species is the most problematic, due to de-
tion) found in the white- and sand-coloured tanks.
pendence on live food (Sykes, Domingues, Correia
Black tanks promoted the lowest and highest values
et al. 2006). Feeding is the most crucial factor deter-
for total mortality and biomass respectively. For
mining both survival and development, and its im-
juveniles, the use of dierent tank colours promoted
portance is enhanced by temperature (Domingues,
dierent growth (Po0.05), but not mortality. Black
Bettencourt & Guerra 2006; Sykes, Domingues & An-
tanks promoted the best results in terms of growth
drade 2006) and competition. Nonetheless, other re-
and biomass. The results obtained in the present
lated eco-biological factors should also be of great
study advise the use of black (or dark colour) tanks
importance to achieve good results. In fact, system
in the hatchling and early juvenile stages to reduce
design may aect the success of culturing cuttlesh
the standard deviations associated with growth,
(Lee, Turk, Forsythe & DiMarco 1998), promoting
mortality and biomass production. This will contri-
stress and lower animal welfare as described by
bute to minimize problems associated with slow and
Conte (2004) for nsh.
fast growers and competition.
In order to enhance growth and survival rates dur-
ing the post-hatching phase of S. ocinalis, a number
Keywords: biomass production, culture stage,
of experiments related to the surrounding environ-
European cuttlesh, Sepia ocinalis, tank colours
ment under culture conditions have already been
conducted by Boletzky (1996), Dickel, Boal and Budel-
mann (2000), Sykes, Domingues, Loyd, Sommereld
Introduction
and Andrade (2003) and Poirier, Chichery and Dickel
The cuttlesh (Sepia ocinalis, Linnaeus) is one of the (2004). Lee et al. (1998) stated that tank colour is
most well-known cephalopods (Boletzky 1983) and a extremely important in determining hatchling survi-
good candidate to become one of the rst cephalopod val in squid, while it is not so important in sepia. In
species in commercial aquaculture (Boucaud-Camou fact, the use of black walls seems to be benecial in
1990; Barnabe 1996). Recently, Sykes, Domingues, terms of stress and well-being to other cephalopod
Correia and Andrade (2006) reviewed the aquacul- 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; ing the rst 40 DAH (fed live prey), and juveniles from
Vidal, Dimarco,Wormuth & Lee 2002). Nabhitabhata 30 to 80 DAH (with a transition of live to frozen prey
and Nilaphat (2000) were pioneers in applying the as part of the commonly used weaning protocol).
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 Material and methods
preference for brightness, they used white, grey and
Experiment I ^ hatchlings
black tanks. Sepia pharaonis individuals showed a
preference for low levels of brightness, which were The experiment lasted for 40 days, corresponding to
associated with rst grey and then black tanks. the hatchling stage. A total of 180 cuttlesh hatchl-
Nevertheless, the same authors also stated that ings with a mean wet weight (MWW) of 0.084 
this preference was highly species dependent, and 0.015 g were used. All cuttlesh hatched in the same
on particular physiological needs, and the study was day, from a single brood obtained from cultured
not growth related and rather choice driven. females reproducing in the facilities of the IFAPA
In sh species, tank colour is used to enhance Agua del Pino Experimental Aquaculture Station
growth and survival, and diminish mean weight (Cartaya, Huelva, Spain). The embryonic develop-
standard deviations under culture conditions. For in- ment of eggs as well as broodstock rearing followed
stance, the reectance and colour of tank walls was the protocols described in Sykes, Domingues and An-
shown to increase the prey capture rates in larval drade (2006).
haddock (Downing & Litvak 1999) and grouper (Dur- A ow-through system composed of nine similar
ay, Estudillo & Alpasan 1996), thus inuencing both rectangular tanks was used. Each tank had a total vo-
growth and survival. Nonetheless, until now, the in- lume of 10 L and a bottom area of 0.141m2. The water
uence of tank colours had not been studied in the ow was set to 17 L h  1. Water was pre-ltered
European cuttlesh, being animals usually reared in through an industrial sand lter, decanted and, be-
white tanks in our culture facilities. fore entering the tanks, passed through a 25 mm mesh
Therefore, the main question is whether tank col- lter and a UV bacteriological lter. Water tempera-
our would somehow inuence cuttlesh growth, the ture, salinity and dissolved oxygen were measured
mean weight standard deviations and mortality, daily in the morning. Both temperature and dissolved
and whether this colour should be black to resemble oxygen were measured using a CRISON OXI45 (Bar-
darkness (night conditions) or whether it should be celona, Spain) probe, while salinity was measured
sandy-like to resemble the natural surrounding con- using a Kikuchi optical salinity meter. The mean cul-
ditions in the wild during daylight. Consequently, the ture temperature, salinity and dissolved oxygen con-
objective of the present study was to determine how ditions are presented in Table 1. A natural daylight
dierent colours (white, sand or black for hatchlings, cycle was provided (14 h L:10 h D), which corre-
and black or white for juvenile) aected the growth, sponded to the local natural conditions in spring. In-
survival, tank biomass production of cuttlesh dur- cident and reective light intensity were measured

Table 1 Experimental conditions for both trials

Tank colour Black Sand White

Experiment I
Temperature (all) 19.3  0.5 1C
Salinity (all) 36.7  0.9 g L  1
Dissolved oxygen (all) 7.4  0.1 mg/L (98.2  1.3%)
Incident light intensity (all) 320.00  30.00 lx
Reflective light intensity 10.63  1.27 lxa 124.03  11.89 lxb 130.77  4.18 lxb
Experiment II
Temperature (all) 20.5  1.0 1C
Salinity (all) 38.2  1.0 g L  1
Dissolved oxygen (all) 7.0  0.4 mg/L (95.0  5.2%)
Incident light intensity (all) 320.00  30.00 lx
Reflective light intensity 3.80  0.62 lxa 46.53  9.00 lxb

Superscript letters represent the grouping of dierences within the same row for Po0.001.

442 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 ment, only the colours producing the higher
Italia, Faenza, Italy), and the mean recorded values are dierence in light reection were used: white and
presented in Table1. Dierences in light reection were black. Also, because white and yellow sandy dis-
only found between the black tanks and the remaining played similar light reection and the rst is com-
two colours (Po0.001). To ensure these light condi- monly used in the rearing of this life stage, white
tions, tanks were cleaned daily by suction with a small was chosen for this second trial.
air hose and with a sponge on sampling days. The A ow-through system composed of six similar
choice of the colours tested was made according to rectangular raceway-type tanks (three white and
the following: white ^ commonly used in the eld sta- three black) was used. This system was similar to that
tion of the Ramalhete (CCMAR, University of the Al- of experiment I, except for the size, shape and depth
garve, Faro, Portugal) for cuttlesh culture in the past of the tanks. Each tank had a total volume of approxi-
10 years; yellow sandy ^ used to resemble a sandy bot- mately 40 L and a bottom area of 0.284 m2. To prevent
tom without all its disadvantages (Sykes et al. 2003); cuttlesh from jumping out, all tanks were covered
and black ^ in order to stimulate prey catching as sug- with a 1mm plastic white mesh.
gested by previous authors (Hanlon & Messenger1988; Water temperature, salinity, dissolved oxygen and
Budelmann, Riese & Bleckmann 1991; Boletzky 1996). light intensities were monitored using the same pro-
All paint used in the present experiment was polyur- cedure as described for experiment I and the re-
ethane plus acrylic enamel based. Ink references were corded values are presented in Table 1. A natural
from the ARALCIN paint line (CIN inks) and as follows: daylight cycle and light intensity at the air^water
black ^ C1 80-301.0531/06; sand ^ C1 80-301.0507/06; tank interface similar to that of experiment I was
and white ^ C180-301.0501/06. provided. Dierences in light reection were found
Newly born cuttlesh hatchlings were used for the between black and white tanks (Po0.001).
three dierent-coloured tanks tested. Each replicate/ Each replicate/tank contained 35 cuttlesh juve-
tank contained 20 hatchlings, thus not representing niles, thus not representing a high density nor in the
a high density nor in the limit of bottom areas estab- limit of bottom areas (Sykes et al. 2003).
lished by Sykes et al. (2003), which generate stress During the rst 20 days of experiment, live mysid
and mortality. shrimp (M. slabberi) was supplied ad libitum. After
Because no grass shrimp was available near the that, frozen adult grass shrimp (Palaemonetes varians)
culture facilities, live mysids (Mesopodopsis slabberi) was supplied as food in a quantied ad libitum. There-
and adult Artemia sp. were collected from ponds and fore, cuttlesh were fed in excess but the amount sup-
salt ponds respectively. Mysids were used as the plied was quantied and based on the remains that
primary feeding prey, while Artemia sp. was only were removed approximately 24 h later, i.e., the next
added when there were not enough mysids to cover morning. The quantication was performed by the
the feeding necessities of all the individuals/tanks. overall cuttlesh weight (total biomass) present in
Cuttlesh were always fed ad libitum and, when each tank, from growth data collected every 10 days.
needed, with the same mysid^artemia proportions.

Data collection and analysis


Experiment II ^ transition from hatchling to
Before the start of each experiment, cuttlesh in each
the juvenile stage
tank colour were weighed and an ANOVA test (Zar
A total of 210 cuttlesh juveniles, 30 days of age, and 1999) was performed to ensure that no signicant
with an MWWof 0.488  0.146 g were used. All cut- dierences were found among replicates. In both ex-
tlesh used were born on the same day and from egg periments, all cuttlesh were weighed every 10 days.
clusters collected in the intertidal zone of the beach Data collected were used to calculate (1) the MWW;
of El Rompido (Cartaya, Huelva, Spain). Technology (2) the mean absolute instantaneous growth rate
and location similar to that described for experiment (MAIGR) (%BWday  1) 5 (LnW2  LnW1)/t  100,
I was used. After hatching and during the rst 30 where W2 and W1 are the nal and the initial mean
DAH, cuttlesh were fed live mysids (M. slabberi) and weight for each colour treatment, respectively, Ln
kept in similar white rectangular tanks. the natural logarithm and t the number of days of
Because no signicant dierences among tank col- the total experiment; (3) the total absolute mortality
ours have been calculated for the previous experi- (TAM), which expresses all dead animals for each col-

r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449 443


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


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
Figure 1 Growth in the mean wet weight (g) of hatchl-
ings of dierent-coloured groups (B, black; S, sand and W, colours at the end of the experiment (Table 3).
white) and exponential growth curves with respective Mortality was only observed in a white tank (TAM:
equations for experiment I (Exp. B, black; Exp. S, dark grey; two individuals ^ 6% for the tank; Table 2) and was
and Exp.W, light grey). Data are presented in oset.Vertical not statistically dierent in terms of MCM (P40.05;
lines represent the standard deviation. 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

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.
Figure 2 Mean cumulative mortality [MCM (%)] of MAIGR is commonly used in cephalopod growth ex-
hatchlings of dierent-coloured groups (B, black; S, sand periments, which is based on mean weights and
and W, white) for experiment I. Data are presented in o- could overlook the eect of mortality. Therefore, we
set.Vertical lines represent the standard deviation.
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


Figure 3 Growth in mean wet weight (g) of juveniles of hatchlings and juvenile cuttlesh
dierent-coloured groups (B, black and W, white) and lin-
ear growth curves with respective equations for experi- We believe that the lack of dierences found in experi-
ment II (Exp. B, black and Exp. W, dark grey). Data are ment I was mainly due to the high variability (higher
presented in oset. Vertical lines represent the standard standard deviation ^ Table 2) found in both white and
deviation. sand tanks. In fact, the use of black tanks in this stage
led to less variable values of mortality and biomass
Dierences were found in B (Po0.05; Table 3), gain. As for experiment II, dierences were found be-
where black tanks attained the higher biomass (Table tween colours for all the indexes used, except for
2). Black tanks displayed a mean B of 106.1  5.0 g, mortality.
while white tanks attained 80.7  7.5 g. Accordingly, Most of the mortality in cuttlesh culture occurs at
B% displayed dierences (Po0.05) between colours the hatchling stage (Sykes, Domingues, Correia et al.
at the end of the experiment (Table 3), and was high- 2006) and this may be caused by several factors al-
er in black tanks (Table 2). ready identied, such as the amount and quality of
prey (Domingues, Kingston, Sykes & Andrade 2001;
Domingues, Sykes & Andrade 2001; Domingues,
Discussion Sykes, Sommereld, Almansa, Lorenzo & Andrade
2004), or minimum bottom areas (Sykes et al. 2003),
A methodological point: biomass production
and others. We intended to understand the eects of
as a new variable
tank colour on cuttlesh growth and survival but,
This study intended to incorporate two culture stages while analysing the results, we realized that those
(hatchling and juvenile) and their transition (which cannot be dissociated from light intensity (which

446 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.

was similar among tanks for incident light but com- to perceive objects in their background that dier in
pletely dierent for reective light). In fact, black contrast by 15%. At the time of feeding, while indivi-
tanks displayed a signicantly lower reective light duals in black tanks remained quiet and camouaged
intensity than the remaining tested colours, and the in the bottom, waiting for the prey to get into their
present results reect this dierence. The present re- visual striking area, white and sand individuals
sults were dierent from those obtained by Nabhitab- would move more in the tanks looking for the prey.
hata and Nilaphat (2000) for S. pharaonis in terms of According to Marshall and Messenger (1996), the
colour. Nonetheless, grey was not tested in the pre- species neural anatomy allows it to camouage with
sent study and neither was tank colour evaluated in the surrounding environment, because it is able to
that study. We believe that a combination of black perceive dierent light intensities in the background
tanks and incident light intensity might promote and generate contrast-dependent patterns. This abil-
adequate brightness and contrast conditions and ity is used by cuttlesh for hunting and concealment
stress the importance of conducting experiments re- (camouage) and has been described extensively by
garding brightness and contrast by using dierent other authors, such as Hanlon and Messenger (1988,
light intensities. This is due to the fact that, according 1996).
to the latter authors, brightness and contrast will in- In experiment I, when the size dierences among
terfere with visual discrimination and, therefore, the individuals increased, larger cuttlesh present in
hunting. 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-
Theoretical relationship between cuttlesh
ably led to even higher dierences in size that
visual acuity and predation behaviour
eventually resulted in starvation and mortality.
While developing culture systems for a given species, Nevertheless, in experiment II, because dead prey
its biology and ecology must be considered. The Eur- remained quiet at the bottom of the tank, cuttlesh
opean cuttlesh is a visual hunter (Messenger 1968) present in the black tanks theoretically had to spend
and has an excellently developed neural anatomy, the same amount of energy as those in white tanks
which is responsible for its distinctive sensory abil- for hunting. Therefore, the dierences found may be
ities, which include mechanoreception, chemorecep- related to the capacity of cuttlesh to see the prey, i.e.
tion and vertebrate-like vision (Budelmann & Young the brightness and contrast of the tanks, which is
1993; Boal, Shashar, Grable,Vaughan, Loew & Hanlon light intensity related.
2004), all of them contributing to prey hunting. The use of dierent tank colours, with a similar in-
Cuttlesh eyes display high sensitivity to light and cident light, produced distinct reective light intensi-
a visual acuity that is far better than that of sh ties in the tanks, thus producing dierent brightness
(Watanuki, Kawamura, Kaneuchi & Inashita 2000), and contrast in the tanks.
which is due to their photosensitive pigments (Wells
1966; Hara 1968). The degree of cuttlesh pupil con-
striction is ambient light intensity dependent (Dou-
Final considerations
glas, Williamson & Wagner 2005) and, during the
present study, reective light in both sand and white Black tanks facilitated easier tank cleaning because
tanks was10 times higher than that in black tanks. In the remains of dead and uneaten prey were more visi-
fact, the retinal structures of cuttlesh are ideal for ble than in the other tanks, especially while using
optimal light capture in dark environments (Davies, grass shrimp. Generally speaking, this colour im-
Gowen, Krebs, Schertler & Saibil 2001) and do not re- proved growth and survival, which were translated
spond well to bright light, as they lack a consensual into higher biomass increase and more accurate and
light-induced pupil response (Douglas et al. 2005). precise growth values. Also, the use of black tanks in
The fact that the species is colour blind (Marshall & the hatchling and juvenile stages can be used as a so-
Messenger 1996; Mthger, Barbosa, Miner & Hanlon lution to reduce the problem with slow and fast
2006) does not interfere with the quality of its vision, growers resulting from competition. Nevertheless,
because adequate visual information is gathered by there is a need for further studies regarding the use
contrasting light and dark shades (Boal et al. 2004). of black tanks with other culture variables such as
According to Mthger et al. (2006), cuttlesh is able at the adult reproducing stage, when an articial diet

r 2010 Blackwell Publishing Ltd, Aquaculture Research, 42, 441^449 447


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 Dickel L., Boal J.G. & Budelmann B.U. (2000) The eect of
stage of the life cycle. Additional studies are also ne- early experience on learning and memory in cuttlesh.
cessary to clarify the inuence of incident and reec- Developmental Psychobiology 36,101^110.
tive light intensity in every culture stage and using Domingues P., Sykes A., Sommereld A., Almansa E., Lorenzo
black tanks. A. & Andrade J.P. (2004) Growth and survival of cuttlesh
(Sepia ocinalis) of dierent ages fed crustaceans and sh.
Eects of frozen and live prey. Aquaculture 229, 239^254.
Domingues P.M., Kingston T., Sykes A. & Andrade J.P. (2001)
Growth of young cuttlesh, Sepia ocinalis (Linnaeus
Acknowledgments
1758) at the upper end of the biological distribution tem-
Antonio Sykes wishes to thank the Fundacao para a perature range. Aquaculture Research 32, 923^930.
Ciencia e a Tecnologia, which funded the current re- Domingues P.M., Sykes A. & Andrade J.P. (2001) The use of
search by way of a PhD (SFRH/BD/12409/2003) and Artemia sp. or mysids as food source for hatchlings of the
a Post-Doc (SFRH/BPD/36100/2007) grant. The rst cuttlesh (Sepia ocinalis L.); eects on growth and survi-
author is also indebted to the IFAPA, from the Junta val throughout the life cycle. Aquaculture International 9,
319^331.
de Andalucia, which allowed him to conduct the ex-
Domingues P.M., Bettencourt V. & Guerra A. (2006) Growth
periments at the CIFAP Agua del Pino Aquaculture
of Sepia ocinalis in captivity and in nature.Vie Et Milieu-
Research Facilities.We also wish to thank two anon-
Life and Environment 56, 109^120.
ymous referees for their valuable criticism. Douglas R.H.,Williamson R. & Wagner H.J. (2005) The pupil-
lary response of cephalopods. Journal of Experimental
Biology 208, 261^265.
Downing G. & Litvak M.K. (1999) The eect of photoperiod,
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