Aquaculture Research, 2005, 36, 1588^1594 doi:10.1111/j.1365-2109.2005.01383.

The impact of light intensity and algal-induced

turbidity on first-feeding Seriola lalandi larvae

Alexander G Carton
Leigh Marine Laboratory, Warkworth, New Zealand

Correspondence: A G Carton, Leigh Marine Laboratory, PO Box 349,Warkworth 1241, New Zealand. E-mail: a.carton@auckland.ac.nz

Abstract tems is paramount as it is these systems that directly

underpin exogenous feeding and thereby play a crucial
Feeding performance (intensity and incidence) of
role in survival (Blaxter 1969, 1986). Because energy
¢rst-feeding yellowtail king¢sh larvae was evaluated
stores are directly proportional to weight, larvae have
under a range of light intensities and algal-induced
little stored energy to withstand periods of starvation
turbidities. Larvae were fed with varying degrees
(Fuiman 2002). Consequently, if exogenous feeding is
of success under all light intensities tested (0^
not initiated shortly after absorption of the yolk sac,
17 mmol s
1 m
2), in both clearwater and green-
larvae reach a ‘point of no return’and fail to feed even
water (8  104 cells mL
1). There was a consistent
if given the opportunity to do so (Yin & Blaxter 1987;
trend for feeding performance to increase with larval
Bisbal & Bengtson 1995). The period between yolk sac
age and light intensity in both clearwater and green-
absorption and the point of no return then represents a
water conditions, demonstrating that visual pro¢-
critical period when ¢rst feeding must be initiated.
ciency increases with larval age. Feeding intensity
In the majority of larvae investigated to date, prey
remained low over the ¢rst 3 days of ¢rst feeding
detection is primarily mediated by the visual system
across all light intensities tested. Days 6 and 7 post-
(Blaxter 1968, 1969; Batty 1987; Browman & O’Brien
hatch larvae showed considerably higher feeding in-
1992; Job & Bellwood 1996). Consequently, the initia-
tensities particularly at 8 and 17 mmol s
1 m
2. This
tion and continuation of exogenous feeding will be
improvement indicates an ontogenetic shift in sen-
most successful if larvae and prey are matched in
sory or locomotor competence. First-feeding larvae
time and space (Cushing 1972), and under conditions
performed equally well in both clearwater and green-
that are most advantageous to visually mediated
water (8  104 cells mL
1) conditions, although at
feeding. Visual feeding thresholds for larval ¢sh are
the lowest light intensity tested (0.1 mmol s
1 m
2)
highly variable across species (Blaxter 1968, 1969;
feeding performance was noticeably reduced. Feed-
Batty 1987; Pankhurst & Hilder 1998; Downing & Lit-
ing performance was severely diminished across
vak 2001; Pena, Dumas, Saldivar-Lucio, Garcia,Tras-
all larval ages at an algal cell density of 32 
vina & Hernandez-Ceballos 2004), an e¡ect that most
104 cells mL
1, demonstrating that for this species
likely results from species-speci¢c di¡erences in ret-
algal-induced turbidities 416  104 cells mL
1 ad-
inal morphology and/or development. Consequently,
versely a¡ect the ability to capture free-swimming
illumination levels that are conducive to ¢rst feeding
prey during the ¢rst-feeding window.
in a larviculture situation will vary according to the
species being cultured.
Keywords: light intensity, greenwater culture,
Larviculture often employs a technique in which
¢rst feeding, feeding performance
turbidity is induced by the addition of algal cells into
the culture medium, a technique commonly referred
Introduction
to as greenwater culture. This technique has been
Many marine teleosts have a reproductive strategy shown to improve survival in larviculture operations
that involves producing numerous small pelagic eggs, (Naas, Naess & Harboe 1992; Gulbrandsen, Lein &
which hatch with a small endogenous energy supply Holmefjord 1996; Lazo, Dinis, Holt, Faulk, & Arnold
in the form of a yolk sac (Fuiman 2002). During this 2000; Eddy & Jones 2002; Papandroulakis, Divanach
period the development of sensory and locomotor sys- & Kentouri 2002), although the precise mechanism

1588 r 2005 Blackwell Publishing Ltd


Aquaculture Research, 2005, 36, 1588^1594
remains unknown. In some species turbidity has
been shown to enhance visually mediated feeding
(Ben¢eld & Minello 1996; Rieger & Summerfelt 1997;
Utne 1997; Cobcroft, Pankhurst, Hart & Battaglene
2001), while in others feeding performance has re-
mained una¡ected (Gardner1981) or declined (Moore
& Moore 1976). Collectively these studies illustrate a
species-speci¢c component in the e¡ect of turbidity
on feeding performance. Often the in£uence of tur-
bidity on larval feeding resembles a dose^response-
type function, with feeding performance declining
at a particular turbidity. Consequently, the level of al-
gal-induced turbidity used in larviculture will have a
profound in£uence on both the initiation of ¢rst feed-
ing and subsequent feeding performance.
Yellowtail king¢sh, otherwise known as Gold-
striped amberjack (Seriola lalandi, Valenciennes), are
a sub-tropical marine ¢sh of the Carangidae family.
The aquaculture of Seriola spp. is well established in
Asia, with sea cage culture of yellowtail (S. quinquer-
adiata) in Japan being one of the largest and most
pro¢table aquaculture industries in the world (Bene-
tti, Nakada, Minemoto, Hutchinson, Shotton & Tin-
dale 2001). The Japanese industry principally depends
on wild caught juveniles, which are then ongrown to
marketable size (Benetti et al. 2001). Because of the
species exceptional growth characteristics, high de-
mand and price on the international market S. lalandi
is currently being assessed as a potential aquaculture
candidate in the Australasian region (Poortenaar,
Hooker & Sharp 2001). The New Zealand culture of S.
lalandi is solely reliant on hatchery production, how-
ever, high mortalities experienced during early devel-
opmental stages are seen as a bottleneck to ¢ngerling
production in this species (Benetti et al. 2001). Poor
feeding performance and low rates of ¢rst feeding in
early larvae appear to indicate that inappropriate cul-
ture conditions during the ¢rst-feeding window may
have contributed to high mortality in initial rearing
trails. The aim of this study was to examine the feed-
ing performance of yellowtail king¢sh larvae at
various light intensities and various algal-induced
turbidities over the ¢rst-feeding period with the speci-
¢c intention of determining optimal larviculture para-
meters that promote the initiation of ¢rst feeding.
Materials and methods
Egg collection and incubation
Broodstock were held at the Bream Bay Aquaculture
Park (National Institute of Water and Atmospheric
r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1588^1594
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E¡ect of light & turbidity on ¢rst-feeding Seriola spp. A G Carton
Sciences, New Zealand), under ambient photoperiod
and water temperature. Fertilized eggs, from natural
spawning events, were collected each morning
(  0700 hours) and stocked into 250 L black cylin-
dro-conical upwelling tanks and incubated at
20.5 1C (range 18.7^21.2 1C). Natural salt water was
¢ltered to 1 mm, UV sterilized and maintained at
0.5 L
1 min
1. Incubation was conducted under
simulated ambient photoperiod, using £uorescent
white light (Philips Alto, Auckland, New Zealand)
producing a light intensity at the surface of
2.8 mmols s
1 m
2. Each day 4 L of water was
drained from the base to remove any dead eggs. Lar-
vae began to hatch after  3 days incubation (ca.
72 h), after which 10 L of water was drained from the
incubator to remove egg debris, and the water £ow
increased to 1L
1 min
1. Following hatching, larvae
remained in the incubators.
Experimental apparatus and lighting
Feeding trials were conducted in black circular
containers (20 cm (D)  10 cm (H)), ¢lled with 2 L
of 1 mm ¢ltered seawater. Containers were lightly
bubbled with air at the center to prevent prey from
settling down and to ensure a homogeneous mix of
algae; this did not noticeably a¡ect larval locomotion
or orientation. White light was generated by Osram
50 decostar bulbs (400^700 nm) (OSRAM Australia,
Sydney, Australia), ¢xed 1.2 m above the tank. Lights
were mounted in the ceiling of a light-tight cabinet
that was divided into ¢ve separate compartments,
each with its own light source. Intensities were ad-
justed via a rheostat and measured at the water sur-
face using a Li-Cor Li-185A light meter (LI-COR
Biosciences, Lincoln, NE, USA). Intensities were
checked prior to and at the conclusion of each trial.
Water temperature within the experimental contain-
ers remained static even under the highest light in-
tensity tested.
Sampling and analysis of gut contents
Following yolk sac absorption, larvae were fed twice
daily (09:00 and 14:00 hours) with rotifers (Brachio-
nus plicatilis enriched DHA Selco, INVE Aquaculture,
Belgium) at 6 mL
1. On the morning of each experi-
ment, feed was withheld and  350 larvae were re-
moved from the rearing tank and transferred to a
20 L holding tank using a ladle. Fifteen larvae were
then sampled from the group to con¢rm gut evacua-
tion prior to experiments. Twenty larvae were then
1589

E¡ect of light & turbidity on ¢rst-feeding Seriola spp. A G Carton
ladled into each of the test containers (10 larvae L
1)
and placed at random into one of ¢ve light-treatment
compartments. Larvae were allowed to acclimatize
for15 min before the introduction of live feed (rotifers).
At the start of each trial containers were stocked with
rotifers at a density of 7 mL
1, the addition of which
was staggered at 8 min intervals across trials, this al-
lowed a su⁄cient amount of time to assess feeding at
the conclusion of each trial. After 2 h larvae were col-
lected on an 800 mm screen, washed with 1 mm ¢l-
tered seawater and transferred on to cold (
2 1C)
microscope slides, a cover slip lowered on top, and ex-
amined under a dissecting microscope. Using this
method gut contents are either expressed or made
clearly visible through the transparent straight gut.
Individual rotifers can then be identi¢ed whole or by
the undigested mastax (Downing & Litvak 2001).
Feeding performance was evaluated as feeding inten-
sity (rotifers consumed per larva per hour) and feed-
ing incidence (proportional of larvae feeding).
E¡ect of light intensity on feeding
performance in clearwater
Experiment1examined the e¡ect of light intensity on
the feeding performance of days 3^7 post-hatch lar-
vae in clearwater conditions. Five light intensities
were tested, 0, 0.1, 1, 8 and 17 mmol s
1 m
2. There
were six replicates for each treatment across two lar-
val cohorts (3 replicates cohort
1).
E¡ect of light intensity on feeding
performance in turbid conditions
Experiment 2 examined the e¡ect of light intensity
on feeding performance of days 3^6 post-hatch lar-
vae in greenwater conditions (8  104 cells mL
1,
Chaetoceros muelleri). Light intensity and level of re-
plication were identical to those used in the clear-
water experiment.
E¡ect of variable algal-induced turbidity on
feeding performance
Experiment 3 examined the e¡ect of increasing levels
of algal-induced turbidity on the feeding intensity
and proportion of larvae feeding in days 3, 4 and 5
post-hatch larvae under supra-threshold light inten-
sity. Four algal densities were used, 0, 8, 16 and
32  104 cells mL
1, light intensity at the water sur-
face for all treatments was set at 12 mmol s
1 m
2.
Low production of eggs prevented assessment of old-
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Aquaculture Research, 2005, 36, 1588^1594
er larvae and a higher level of replication. There were
three replicates for each treatment across a single lar-
val cohort.
Statistical analysis
To standardize development larval age was converted
to degree days prior to statistical analysis. All data
were tested for normality (Kolmogorov^Smirnov
test) and homogeneity of variance (Fmax test). To ac-
complish this, proportional data (feeding incidence)
were arcsine transformed and feeding intensity data
logarithmically (x 0 5 log (x11)) transformed. A one-
way ANOVA was then used to analyse the e¡ect of light
intensity or algal-induced turbidity treatments on the
feeding intensity and proportion of larvae feeding at
each age. When treatment e¡ects were signi¢cant, a
Holm^Sidak multiple comparison test was employed.
Data are reported as mean values  SEM.
Results
E¡ect of light intensity on feeding incidence
in clearwater conditions
The incidence of ¢rst feeding at day 3 post-hatch was
signi¢cantly di¡erent among treatments (Po0.001),
peaking at the two highest light regimes tested
(Fig. 1a). This remained consistent in day 4
(Po0.001) and day 5 (Po0.001) post-hatch larvae.
Large increases in feeding incidence under 8 and
17 mmol s
1 m
2 were clearly evident between 3
and 4 days post hatch (DPH), increasing from
30.1  4.6% to 56.7  10.3% under the highest light
intensity tested. Feeding incidence was signi¢cantly
di¡erent among treatments on day 6 (P 5 0.001) and
day 7 (P40.001), although feeding incidence at
1 mmol s
1 m
2 was not signi¢cantly di¡erent from
that observed under 8 or 17 mmol s
1 m
2. Light in-
tensity also had a signi¢cant e¡ect on feeding inten-
sity at all larval ages. In general, feeding intensity
increased with both larval age and light intensity
(Fig. 1b). Feeding intensity of day 3 post-hatch larvae
was low although it di¡ered signi¢cantly among
treatments (Po0.001). At 4 DPH feeding intensity
progressively increased with light intensity, peaking
at the two highest light regimes tested (Po0.001).
Feeding intensity of days 5^7 post-hatch larvae
peaked at 8^17 mmol s
1 m
2, although feeding in-
tensity under these light regimes was not signi¢cantly
di¡erent from that observed under 1 mmol s
1 m
2.
r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1588^1594
 13652109, 2005, 16, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2109.2005.01383.x by INIDEP-Instituto Nacional de Investigacion y Desarrollo Pesq, Wiley Online Library on [05/07/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Aquaculture Research, 2005, 36, 1588^1594 E¡ect of light & turbidity on ¢rst-feeding Seriola spp. A G Carton

(a) 90 (a) 90
80 0 80 0
0.1 0.1

Feeding incidence (%)

Feeding incidence (%)
70 70
1 1
60 8 60 8
17 50 17
50
40 40

30 30

20 20

10 10

0 0
3 4 5 6 7 3 4 5 6
Age (days post hatch) Age (days post-hatch)

(b) 6 (b) 6

5 5
Feeding intensity

Feeding intensity
4 4

3 3

2 2

1 1

0 0
3 4 5 6 7 3 4 5 6
Age (days post hatch) Age (days post-hatch)

Figure 1 (a) Feeding incidence (mean  SEM) and (b)
feeding intensity of ¢rst-feeding yellowtail king¢sh larvae in
clearwater at ¢ve di¡erent light intensities (0, 0.1, 1, 8 and
17 mmol s
1 m
2). N 5 6 replicates, 20 larvae per replicate.
Di¡erent letters within larval ages indicate signi¢cant di¡er-
ences among light intensity treatments (Po0.05). Feeding in-
tensity is measured as rotifers consumed per larva per hour.
Feeding performance under total darkness was ap-
preciably impaired across all larval ages.
E¡ect of di¡erent light intensities and single
level of algal-induced turbidity
Feeding performance in greenwater conditions
showed similar trends as those observed in clear-
water with both feeding incidence and intensity
increasing with light intensity and larval age
(Fig. 2a and b). Algal-induced turbidity of 8 
104 cells mL
1 severely reduced both feeding inci-
dence and intensity of days 5 and 6 post-hatch larvae
in the 0.1 mmol s
1 m
2 treatment.
Figure 2 (a) Feeding incidence (mean  SEM) and (b)
feeding intensity of ¢rst-feeding yellowtail king¢sh larvae
in greenwater (8  104 cells mL
1) at ¢ve di¡erent light
intensities (0, 0.1, 1, 8 and 17 mmol s
1 m
2). N 5 6 repli-
cates, 20 larvae per replicate. Di¡erent letters within lar-
val ages indicate signi¢cant di¡erences among light
intensity treatments (Po0.05). Feeding intensity is mea-
sured as rotifers consumed per larva per hour.
cell density (P 5 0.422). Feeding incidence in the
32  104 cells mL
1 treatment was signi¢cantly dif-
ferent from all other treatments in day 4 (P 5 0.004)
and day 5 (P 5 0.002) post-hatch larvae, with feeding
incidence being consistently higher at 0, 8 and
16  104 cells mL
1 treatments (Fig. 3a and b). In
the case of days 4 and 5 post-hatch larvae both the
incidence and intensity of larval feeding was drasti-
cally depressed at the highest algal cell density tested
(32  104 cells mL
1).
Discussion

Yellowtail king¢sh larvae consumed prey with vary-

E¡ect of variable levels of algal-induced
turbidity at a single light regime
The incidence of ¢rst feeding was highly variable at
day 3 post hatch and this was independent of algal
ing degrees of success under all light intensities (0^
17 mmol s
1 m
2) tested. Despite this, feeding inci-
dence and feeding intensity in clearwater conditions
were severely depressed in total darkness, indicating

r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1588^1594 1591


E¡ect of light & turbidity on ¢rst-feeding Seriola spp. A G Carton
(a) 90
0 8 16 32
80
70
Feeding incidence (%)
60
50
40
30
20
10
0 in visual capabilities with larval age. Increasing feed-
3 4
Age (days post-hatch)
(b) 2 tent with other studies (Pankhurst & Hilder 1998;
1.5
Feeding intensity
1 of the visual system, locomotor system or a combina-
0.5
0 cess of 50%. This suggests that feeding over this early
3 4
Age (days post-hatch)
Figure 3 (a) Feeding incidence (mean  SEM) and (b)
feeding intensity of ¢rst-feeding yellowtail king¢sh larvae
at four di¡erent algal densities (0, 8, 16 and 32 
104 cells mL
1). N 5 3 replicates, 20 larvae per replicate.
Di¡erent letters within larval ages indicate signi¢cant dif-
ferences among treatments (Po0.05). Feeding intensity is
measured as rotifers consumed per larva per hour.
that yellowtail king¢sh larvae are primarily depen-
dent on vision for feeding. The ability to feed in com-
plete darkness is most likely to be mediated by non-
visual sensory systems, in particular the lateral line
system (Montgomery, Macdonald & Housley 1988;
Jones & Janssen 1992; Janssen, Jones,Whang & Oshel
1995). At hatching, yellowtail king¢sh larvae possess
a number of super¢cial neuromasts distributed over
the head and trunk, and these receptors proliferate
during early development (pers. obs.).
Light intensity may in£uence feeding performance
either by operating directly on the visual system or
indirectly on non-sensory attributes. An increase in
feeding performance with increasing light intensity
has been demonstrated for £at¢sh (Blaxter1969), her-
ring (Blaxter 1968), striped trumpeter (Pankhurst &
Hilder 1998; Cobcroft et al. 2001) haddock (Downing
& Litvak 2001) and spotted sand bass (Pena et al.
2004) larvae. Increase in feeding performance with
increasing light intensity may not solely be because
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Aquaculture Research, 2005, 36, 1588^1594
of enhanced visual detection of prey. For example, ac-
tively feeding herring alter swimming behaviour and
speed in response to variations in light intensity
(Batty 1987). Increases in feeding performance could
then also be attributed to modi¢ed swimming beha-
viour, which may alter the prey encounter rate or
total search volume, or a combination of both.
Feeding performance in clearwater conditions
increased with larval age across all light regimes
(0.1^17 mmol s
1 m
2), suggesting an improvement
5
ing performance with larval development is consis-
Cobcroft et al. 2001; Downing & Litvak 2001). Days 6
and 7 post-hatch larvae showed considerable im-
provement in feeding performance at light intensities
 1 mmol s
1 m
2 indicating that the development
tion of both rapidly improves over this period. Feed-
ing intensity following yolk sac absorption initially
remained relatively low (ca. one rotifer per larva per
hour), although the incidence of feeding was in ex-
5
post-hatch period may be constrained by the develop-
mental state of the visual or locomotor system. Feed-
ing intensity rapidly increased at days 6 and 7 post
hatch, which appears to indicate an ontogenetic shift
in sensory or locomotor competence, or a combina-
tion of both. Improvements in visual sensitivity over
this period may result from an initial increase in cone
density, summation of photoreceptors to gangilion
cells and an increase in eye size and lens diameter,
which collectively improve acuity, sensitivity and vi-
sual range (Higgs & Fuiman 1998a,b).
In general larval feeding performance was com-
parable under clearwater and greenwater conditions;
consequently, algal-induced turbidity appears to
have little positive bene¢t in terms of feeding perfor-
mance. Miner and Stein (1993) found reduced prey
consumption by larval bluegills in turbid conditions
under low (o450 lx)-surface light conditions, but
not at high-surface light conditions. This also agrees
with the observations of Naas, Huse and Iglesias
(1996), demonstrating that in black-walled tanks, as
those used in this study, the addition of greenwater
(25  104 cells mL
1) decreases illumination to-
wards the centre of the tank throughout the water
column, thereby decreasing the amount of light
available for vision.
Varying levels of algal-induced turbidity had a
clear e¡ect on feeding intensity and incidence for
days 4 and 5 post-hatch larvae. At 3 DPH there was
r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1588^1594

Aquaculture Research, 2005, 36, 1588^1594
no di¡erence in feeding performance across the four
algal cell densities tested. However, at days 4 and 5
post hatch, both feeding intensity and incidence were
adversely a¡ected at the highest algal cell density
(32  104 cells mL
1) tested. Reduced feeding per-
formance at the highest cell density may be the result
of increased light attenuation, elevated scattering re-
ducing the ability of larvae to discriminate prey or
changes in the ratio of downwelling to upwelling
light (Naas et al. 1996; Cobcroft et al. 2001).
In contrast, feeding performance at the remaining
algal cell densities was approximately similar, al-
though feeding intensity and incidence peaked at
8  104 cells mL
1 for day 5 post-hatch larvae. These
results demonstrate that for yellowtail king¢sh lar-
vae algal cell densities of 416  104 cells mL
1 have
an adverse impact on the ability to capture free-
swimming prey during the ¢rst-feeding window.
Consequently for this species the optimal greenwater
culture density appears to be between 8 and
16  104 cells mL
1.
The present study demonstrates that the feeding
performance (incidence and intensity) of yellowtail
king¢sh larvae is in£uenced by light intensity and al-
gal-induced turbidity. Greenwater-induced turbidity
does not appear to convey a substantial feeding ad-
vantage with feeding incidence similar in both clear-
water and greenwater conditions, although at day 5
post-hatch feeding intensity in 8  104 cells mL
1
was signi¢cantly higher than other treatments. How-
ever, algal cell densities 416  104 cells mL
1 ad-
versely impact on the ability to capture free-
swimming prey during the ¢rst-feeding window.
These results can assist in developing larviculture
protocols that maximize larval survival by providing
lighting and greenwater culture regimes that aid the
rapid initiation of ¢rst feeding and subsequent transi-
tion through the ¢rst-feeding window.
Acknowledgments
I would like to thank Dr Brendan Gara (National Insti-
tute of Aquatic and Atmospheric Sciences, New Zeal-
and), Damien Moran and Cea Kapiri-Smith (Institute
of Aquatic and Atmospheric Sciences, New Zealand)
for assistance with egg collection, larval rearing and
logistical support.
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