Advantages of LED Lighting Systems

in Larval Fish Culture

Juliette Delabbio

ish need light to

on visual photoreception to
survive and grow; it is one
establish the location of prey
of the essential inputs to
and poor lighting can cause
life. In aquaculture there are
abnormal eye development,
natural and artificial sources
resulting in decreased visual
of light. Regardless of source,
acuity (Rahmann et al. 1979,
environmental lighting in an
Zeutsius and Rahmann 1984).
aquaculture facility drives
Therefore, most fish larvae
key biological processes that
need a minimal threshold of
FIGURE 1. Research on the effects of ONCE LED lighting on growth
influence performance and
light intensity to survive, which
performance of red drum larvae is currently underway at the University of
ultimately affect production
has evolved in relationship to
Texas Marine Science Institute. Photo by Cypress Hansen.
and profitability. Incandescent
the ability to localize, catch
and fluorescent lighting are traditional artificial sources for
and ingest prey in its natural habitat. Larval-rearing environments
environmental lighting, but LED lighting now offers another
in hatcheries usually use artificial lighting conditions that are
option for hatchery managers.
quite dissimilar to the larvaes natural environment. Teleost larvae
Applications of LED lighting in aquaculture facilities have
need a minimum light intensity of 0.1 lux to locate prey properly
only recently become economically feasible, with the cost of lamps (Blaxter 1986). This is a very general statement; later research has
offset by accrued energy savings. More importantly, traditional
indicated that, under aquaculture conditions, finding the correct
light sources, such as incandescent or fluorescent lamps, do not
environmental lighting for marine fish larvae is a complex issue
provide the opportunity to change light spectrum and intensity
with several complementary factors to consider.
and do not provide optimum lighting conditions for larval
culture. Newly-developed LED lighting systems provide flexible
A Complex Issue: Environmental Lighting
adjustment of light intensity, photoperiod and spectrum in a single Requirements Differ with Species
light source, thus serving as a powerful tool to influence growth
Pea et al. (2004) studied the effect of environmental
and survival in larval fish culture.
lighting on the first feeding stage of spotted sand bass Paralabrax
maculatofasciatus. With a fluorescent lighting source, prey capture
The Significance of Environmental Light
increased with light intensity. Light intensities of 0, 100, 400 and
to Larval Production
700 lux were measured at the air-water interface and capture
Light intensity, spectrum and photoperiod have a significant
success was assessed by prey content in individual digestive tracts.
effect on fish at all life stages (Boeuf and Le Bail 1999, Ruchin
Bass larvae were more successful at capturing prey at 400 and
2004, Han et al. 2005, Marchesan et al. 2005). The larval stage
700 lux than at 0 and 100 lux. Because there was no statistical
of any fish species is exceptionally sensitive to environmental
difference in prey capture between 400 and 700 lux, it was
conditions and newly hatched animals are particularly fragile.
postulated that there may be threshold levels of light intensity to
Light receptivity in larval fish changes during development and
activate feeding behavior but, once those thresholds are reached,
this affects feeding behavior, feed intake and survival (Fig. 1).
increases in light levels do not increase feeding activity until an
The effects of light on marine fish larvae have been studied
upper threshold negative response occurs. This research clearly
in the laboratory (Barahona-Fernendes 1979, Puvanendran and
showed that lighting in a specific intensity range was a factor
Brown 1998, Downing and Litvak 1999a and b, Trippel and Neil
in first-feeding activity. Inasmuch as high larval mortality is a
2002, Migaud et al. 2008, Yoon et al 2010, Vollset et al. 2011) and common occurrence in aquaculture, there is a need to establish
the field (Blaxter 1966, 1968, Suthers and Sudby 1996, Gibson
threshold light levels for larval stages of different commercially
et al. 1998). In total, this research indicates that environmental
produced fish species. Compared to conventional lighting sources,
lighting is an important and complex parameter with broad effects LED lights can smoothly reduce intensity from 100 to 0 percent
on growth and survival of marine fish larvae.
without affecting spectral output.
In the laboratory, an increase in light intensity coincided with
The Effect of Light Intensity on
greater success rates of prey capture by haddock larvae (Downing
Marine Fish Larvae Performance
and Litvak 1999a). Atlantic cod Gadus morhua larvae held at high
The larvae of many marine fish species are born predators,
light intensity (680 lux) had lower feeding intensity than larvae
suggesting that the intensity of light in the rearing environment can held at low light intensity (8.5 lux) (Puvanedran and Brown 1998).
(CONTINUED ON PAGE 26)
significantly affect survival and growth. Marine fish larvae rely
W W W.WA S .O R G

W O R L D AQ UACU LT U R E

S E P T E M B E R 2 015

25

Increasing light intensity

increased growth of European
seabass, but there was an upper
limit where light intensity had
a negative effect (BarahonaFernandes 1979). The larvae
of different fish species may
have different threshold levels
of environmental light intensity
(both high and low) and lack of
knowledge of these thresholds
can affect growth and survival.

to change light intensity is

important at the larval stage
after hatch but before first
feeding. Dimmable LED
lighting systems allow the
adjustment of light intensity at
this stage to support optimum
first-feeding activity (Fig. 2).

Geographic Variation
in Response to Light

Most current,
commercially-produced
Timelines on Eye
fish species are not wholly
Structure Development
domesticated. Therefore,
FIGURE 2. Experimental larval system at Mote Aquaculture Research Park
Environmental lighting
when devising an optimum
used to evaluate ONCE Innovations LED lamps and fluorescent lamp on light
needs change during larval
lighting treatment for marine
spectrum and intensity effects.
development, further
fish larvae, important
complicating the choice of
consideration should be
optimum light intensity ranges for larval fish rearing. In marine
given to the specific conditions of the habitat of a particular
fish larvae, the structure of the eye is different at the beginning of
species. Most marine fish species live in a variety of habitats/
the larval stage than at the end (Lyall 1957, Blaxter and Staines
environments during their life cycle and have evolved to adapt to
1970, Blaxter 1975, Hairston et al. 1982, Raymond 1985). Rods are the specific physical characteristics of their environment. Light
not part of the retina of newly hatched larvae. These specialized
intensity and photoperiod at different latitudes might have an
receptors, needed for vision in low light intensity environments,
effect on larval growth and survival (Suthers and Sundby 1996).
develop in the retina as larvae grow. First-feeding larvae, therefore,
Differences in population performance of larval cod are related to
are strictly dependent on cone vision for prey identification and
environmental light intensity (Puvanendran and Brown 1998). In
capture. Cone vision requires high levels of light intensity for
that study, cod larvae from two geographically distinct locations
photostimulation to occur. Consequently a specific light intensity in spawned at different times during the year. The larvae of one
the environment can affect feed intake quite differently, depending
population experienced much higher light intensities in their natural
on the age/stage of larvae.
environment than the other. Cod larvae from the populations had
As larval fish body size increases, visual acuity and its reacdifferent feeding behavior related to environmental light exposure.
tive distance to prey also increases (Blaxter 1986). Consequently,
Application of the natural light conditions of the geographical
as larvae grow larger, they are also more successful at prey capture. environment of origin affected larval culture performance.
In contrast, cod larvae reared at low light intensities during larval
Responsiveness of a species at the population level has been
development had overall lower mortality rates than larvae held at
reported for other environmental parameters (Berg and Moen
higher light intensities (Puvanendran and Brown 2002). There may 1999, Jensen et al. 2000, Wilds and Muoneke 2001, Imsland et al.
be critical junctures where a light intensity that was beneficial previ- 2005, Conover et al. 2009, Burt et al. 2011) but the contribution
ously is no longer applicable or not as beneficial because of develand effect of environmental light on larval survival of different
opmental changes (i.e. improved visual acuity) as the eye structure
populations remains largely unexplored.
develops through larval development, and even through many juvenile stages. Varying light intensity levels for different stages of larThe Significance of Tank Color
val development is easily accomplished with LED lighting systems. to Larval Performance
Environmental light intensity may improve capture success
Prior to First Feeding
because it increases the contrast between prey and background
Light levels for marine fish larvae are important even before
(Pea et al. 2004). There is a broad variation in the color of larval
the start of first feeding. Herring Clupea harengus larvae at the
rearing units and, for larval vision, the contrast of prey to backsac-fry stage (prior to first-feeding) have lower activity levels
ground is variable depending on tank color. Studies have explored
when exposed to high light intensities (Batty 1987). This decrease
the effect of tank background color on larval performance (Martinin activity affects yolk consumption rate and the time to firstRobichaud and Peterson 1998, Downing and Litvak 1999b, Tamfeeding, thereby affecting the proper time for introduction of prey
azouzt et al. 2000, Bransden et al. 2005, Monk et al. 2008, Jirsa et
organisms.
al. 2009). Unfortunately these studies have not proven whether the
Light intensity prior to first-feeding in larvae subsequently
contributing factor(s) to better larval growth was tank color itself,
affects the survival and timing of first feeding. Atlantic halibut
the contrasting effect provided, or a combination of environmental
Hippoglossus hippoglossus yolk-sac fry developed abnormally in
light intensity and light spectrum with the color characteristics of
the presence of light and high mortality subsequently occurred at
the specific prey organism and tank color. Tank color should be a
first feeding (Bolla and Holmefjord 1988). Therefore, the ability
consideration when establishing environmental lighting conditions.

26

S E P T E M B E R 2 015

W O R L D AQ UACU LT U R E

W W W.WA S .O R G

With LED lighting systems, it is possible to change light intensity

and light spectrum in a single light source, and hatchery managers
can establish the best light intensity and spectrum for fish reared
in tanks with a specific background color. This is site specific but
LED lighting allows this kind of customization in environmental
lighting.

Environmental Light and Photoperiod

Photoperiod is the duration of light exposure during a 24-hour

cycle and has an important effect on the growth and survival of
marine fish larvae. Larval fish often change in response to light
as they develop. Survival of early-stage cod larvae raised with
continuous (24-hour) light exposure was significantly greater
than that of larvae reared with a 12- or 18-hour photoperiod
(Puvanendran and Brown 2002). Initially larvae subjected to
continuous light exposure were larger than 12- and 18-hour
photoperiod but, after a certain stage of development, photoperiod
did not affect growth or survival.
For some species, longer photoperiod in early development
stages may create more feeding opportunities, thereby increasing
the likelihood that larvae begin exogenous feeding and have a
greater success rate of prey capture. Young larvae are particularly
fragile, with few internal resources for sustenance if they do not
successfully transition to exogenous feeding. Early high success
rate of prey capture readily translates to better growth, nutrition
and survival. Longer photoperiods often stimulate increased motor
activity and increases in muscle development and agility, although
this can have a negative effect on growth.
However, continuous light exposure during early larval
development is not recommended for all marine fish larvae. There
is a varyiable response to photoperiod length among larvae of
different fish species. The first-feeding stage of rabbitfish Siganus
guttatus had excellent growth under continuous light exposure
(Duray and Kohno 1988). The larvae of black porgy Mylio
macrocephalus had the best survival when reared with a 13-hour
photoperiod (Kiyono and Hirano 1981). Growth of European
seabass Dicentrarchus labrax larvae was greatest with an 18hour photoperiod (Barahona-Fernandes 1979). For black porgy
and European seabass, larvae exposed to continuous light did not
perform as well as larvae reared under photoperiod regimes that
were more reflective of natural habitats.
Additionally there seems to be an intra-species component
to the effect of photoperiod on growth of marine larvae, similar
to findings on intra-species response to light intensity, that must
also be considered in establishing light exposure regimes for larval fish. Under the same photoperiod regime, cod larvae from the
southern range of the species (Scotian shelf region) have a lower
growth rate than larvae from the more northern range (Arctic areas)
(Suthers and Sundby 1996). Northern cod populations have a much
longer summer photoperiod in their natural habitat and, therefore,
are more active for longer periods of time and had a greater prey
capture success rate. LED lighting systems with programmable
controllers allow aquaculturists to specify photoperiod and light
intensity levels within lighting blocks. It is no longer necessary for
photoperiod exposure to be a matter of simple lights-on/lights-off
control.

Effects of Light Spectrum

Light spectrum (color) is one of the most important but most

neglected components of a lighting treatment. Current research
on the effects of light spectrum on performance of marine larvae
is fragmented and difficult to interpret because light treatments
are not well described with respect to the lighting source used and
the method and type of meter used to measure the light spectrum.
At present, there is no standardized method for describing light
treatments. Physical measurements of light spectrum are made
at different places in the rearing environment (i.e. at the air/water
interface, at certain water depths or at the tank bottom) and with
light measurement tools with highly varying sensitivity because of
spectral filters used. Thus, reports on benefits of certain lighting
conditions are not easily replicated and overall prescriptive
statements are severely limited in their application. Nevertheless,
examination of past research provides insights into the effect of light
spectrum on the success and performance of marine larvae.
In fish, the light spectrum is detected by the eye and pineal
gland (Levin and McNicol 1982, Ekstrm and Meissl 1997). Light
spectrum is important at a very early age of development. Similar to
light intensity, light spectrum may be important for successful prey
capture, enabling larval fish to better see the prey through contrast
with the surrounding environment. This is particularly important
during the earliest stages of larval growth, when visual acuity is still
developing.
Because water is a natural filter of light, the light spectrum
received by larvae changes with depth. Larval position in the water
column can mean different spectrum experience and different
contrast abilities, even within a single rearing unit.
The influence of colored light on growth rate of larval Crucian
carp Carassius carassius, rotans Perccottus glenii and guppies
Poecilia reticulata was evaluated by Ruchin (2004). All larvae were
from a single body of water but the fish occupy different ecological
niches and have different feeding behaviors and prey items. The
light intensity at the water surface was the same for each species, but
the color of light in individual niches was quite different. In ponds,
guppies live in the upper layers, rotans live in the middle layers and
carp live in the benthic layers. There are differences in response to
light in different zones experienced by different species of fish.
Using a control lamp, filters were used to subject the fish to
different peak color wavelengths. All species performed best with
exposure to blue and green wavelengths. Red light had a pronounced
negative effect on growth: a 10 percent decrease in guppies, a 9
percent decrease in rotan and a 33 percent decrease in carp. Yellow
light had a severe negative impact (21 percent growth reduction) on
rotan larvae, but was not as significant with guppies or carp. Guppy
performed best with blue light, rotan with blue and green light
and carp with green light. Carp mainly feed on benthic organisms
and rotan, which lives among thickets of vegetation, feed on
zooplankton. Therefore, differences in growth rates of the two fish
species may be related directly to the different degrees of contrast
between background and prey present in the different microhabitats.
The study clearly indicates the importance of light spectrum as
an ecological parameter that can significantly affect larval culture
performance.
Unlike traditional light sources, LED lights can be built to
(CONTINUED ON PAGE 28)

W W W.WA S .O R G

W O R L D AQ UACU LT U R E

S E P T E M B E R 2 015

27

MLA-WH 120V

MLA-RE 120V

MLA-BL 120V

MLA-GY 230V

MLA-WH 120V AT 5%

MLA-RE 120V AT 50%

MLA-BL 120V AT 50%

MLA-GY 230V AT 40%

FIGURE 3. First row shows the spectral output at 100 percent intensity of four LED lamps (ONCE innovations Inc). The second row shows the spectral output
of the same lamps when dimmed to the light intensity levels indicated.

provide a variety of different light color outputs. Figure 3 indicates may have on larval performance. Consequently LED technology
the spectral output of four LED lamps for aquaculture at 100
provides an opportunity to enhance larval fish production.
percent intensity. The light spectra of each lamp would be perceived
as white light but each lamp would have enhanced intensity in a
certain span of wavelengths (color) for a certain stage and/or species Notes
of fish that produces a favorable response in feeding activity. It
Dr. Juliette Delabbio is Director of Research and Development
is also possible to coordinate use of a LED light with particular
for ONCE Innovations Inc., a lighting company specializing in
wavelength specifications with dimming possibilities so that the
LED lighting for aquaculture. Her contact email is jdelabbio@
lighting system can change with the changing needs of fish larvae.
onceinnovations.com.
Figure 3 also indicates the spectral change of lamps when dimming
occurs.

References

Summary

The production of high numbers of robust fish larvae and

subsequent healthy juveniles represent a significant bottleneck in
many marine fish hatcheries. The first-feeding period of all fish
larvae, in nature and controlled culture conditions, is a critical time.
Either larvae are able to identify and aptly respond to food sources
in their environment by successfully capturing and ingesting
enough to remain nutritionally fit or fail to consume sufficient
amounts of food and subsequently die of starvation. Furthermore,
pelagic larvae rely on vision to capture prey and use photoreception
as a means of determining their position in the water column.
In hatcheries, finding the correct environmental lighting for
marine fish larvae is complex, but is an important consideration
because survival of 30 percent is not uncommon in larval fish
culture. Larvae respond to light in their environment and all
three characteristics of artificial environmental light (intensity,
photoperiod and wavelength) contribute to successful larval rearing.
Furthermore, the required timing and modulation of light intensity
and wavelength spectra during larval stages varies with species.
New developments in artificial lighting using LED technology
are providing hatchery managers with a broader range of control
than offered by traditional lighting sources. These new technologies
allow more precise manipulation and measurement of light intensity
and spectrum from a single lighting system and enable aquaculturists to isolate, remove or enhance the effects that these parameters

28

S E P T E M B E R 2 015

W O R L D AQ UACU LT U R E

Barahona-Fernandes, M. 1979. Some effects of light intensity and

photoperiod on the sea bass larvae (Dicentrarchus labrax (L.))
reared at the Centre Oceanologique de Bretagne. Aquaculture
17:311-321.
Batty, R. 1987. Effect of light intensity on activity and foodsearching of larval herring, Clupea harengus: a laboratory study.
Marine Biology 94:323-327.
Berg, O.K. and V. Moen. 1999. Inter- and intrapopulation variation in
temperature sum requirements at hatching in Norwegian Atlantic
salmon. Journal of Fish Biology 54: 636-647.
Blaxter, J.H.S. 1986. Development of sense organs and behaviour
of teleost larvae with special reference to feeding and predator
avoidance. Annual Larval Fish Conference 115:98-114.
Blaxter, J.H.S. 1975. The eyes of larval fish. Vision in Fish. Ed. M.
Ali, NATO Advanced Study Institutes Series 1:427-443.
Blaxter, J.H.S. 1968. Light intensity, vision, and feeding in young
plaice. Journal of Experimental Marine Biology and Ecology
2:293-307.
Blaxter, J.H.S. 1966. The effects of light intensity on the feeding
ecology of herring. Pages 393-409 In: R. Bainbridge, G.C.
Evans, and O. Rackham, editors. Light as an Ecological Factor.
Blackwell, Oxford, England.
Blaxter, J.H.S. and M. Staines. 1970. Pure-cone retinae and
retinomotor responses in larval teleosts. Journal of the Marine
Biological Association of the UK 50:449-460.

W W W.WA S .O R G

Boeuf, G. and P.Y. Le Bail. 1999. Does light have an influence on

fish growth? Aquaculture 177:129-152.
Bolla, S., and I. Holmefjord. 1988. Effect of temperature and light
on development of Atlantic halibut larvae. Aquaculture 74:355358.
Bransden, M.P., T.G.M. Butterfield, J. Walden, L.A. McEvoy and
J.G. Bell. 2005. Tank colour and dietary arachidonic acid affects
pigmentation, eicosanoid production and tissue fatty acid profile
of larval Atlantic cod (Gadus morhua). Aquaculture 250:328-340.
Burt, J.M., S.G. Hinch and D.A. Patterson. 2011. The importance
of parentage in assessing temperature effects on fish early life
history: a review of the scientific literature. Reviews in Fish
Biology and Fisheries 21:377-406.
Conover, D., Duffy, T.A. and L. Hice. 2009. The covariance between
genetic and environmental influences across ecological gradients.
Annals of the New York Academy of Sciences 1168: 100-129.
Downing, G., and M. Litvak. 1999a. The influence of light intensity
on growth of larval haddock. North American Journal of
Aquaculture 61:135-140.
Downing, G. and M.K. Litvak. 1999b. The effect of photoperiod,
tank colour and light intensity on growth of larval haddock.
Aquaculture International 7:369-382.
Duray, M. and H. Kohno. 1988. Effects of continuous light in on
growth and survival of first-feeding larval rabbitfish, Siganus
guttatus. Aquaculture 72:73-79.
Ekstrm, P. and H. Meissl. 1997. The pineal gland of teleost fishes.
Reviews in Fish Biology and Fisheries 7:199-284.
Gibson, R.N., L. Pihl, M. T. Burrows, J. Modin, H. Wennhage
and L.A. Nickell. 1998. Diel movements of juvenile plaice,
Pleuronectes platessa, in relation to predators, competitors, food
availability and abiotic factors on a microtidal nursey ground.
Marine Ecology Progress Series 165:145-159.
Hairston, N.G, K.T. Li and S.S. Easter. 1982. Fish vision and the
detection of planktonic prey. Science 218:1240-1242.
Han, D., S. Xie, L. Lei, Z. Zhu and X. Yang. 2004. Effect of light
intensity on growth, survival and skin color of juvenile Chinese
longsnout catfish (Leiocassis longirostris Gnther). Aquaculture
248:299-306.
Imsland, A.K., A. Foss, W.W. Bonga, E. van Ham and S.O
Stefansson. 2005. Comparison of growth and RNA: DNA ratios
in three populations of juvenile turbot reared at two salinities.
Journal of Fish Biology 60:288-300.
Jensen, A.J., T. Forseth and B. Johnsen. 2000. Latitudinal variation
in growth of young brown trout Salmo trutta. Journal of Animal
Ecology 69:1010-1020.
Jirsa, D., M. Drawbridge and K. Stuart. 2009. The effects of tank
color and light intensity on growth, survival, and stress tolerance
of white seabass, Atractoscion nobilis, larvae. Journal of the
World Aquaculture Society 40:702-709.
Kiyono, M. and R. Hirano. 1981. Effects of light on feeding and
growth of black porgy (Mylio macrocephalus Basilewsky), post
larvae and juveniles. Conseil Permanent International pour
Exploration de la Mer 178: 334-336.
Levin, J. and E. McNichol. 1982. Color vision in fish. Scientific
American 246:108-117.
Lyall, A.H. 1957. The growth of the trout retina. Quarterly Journal
Microscopical Science 98: 101-110.

Marchesan, M., M. Spoto, L. Verginella and E.A. Ferrero. 2005.

Behavioral effects of artificial light on fish species of commercial
interest. Fisheries Research 73:171-185.
Martin-Robichaud, D. J. and R.H. Peterson. 1998. Effects of light
intensity, tank colour and photoperiod on swimbladder inflation
success in larval striped bass, Morone saxatilis Walbaum.
Aquaculture Research 29:539-547.
Migaud, H., A. Davie, S. Carvoni, J. Murray, P.A. Lysaa and J.
Treasurer. 2008. Effects of wavelength on Atlantic cod (Gadus
morhua) larvae performance. Larvi 2009: Fish & Shellfish
Larviculture Symposium. Ghent, Belgium, Sept. 7-9, 2009.
Monk, J., V. Puvanendran and J. Brown. 2008. Does different tank
bottom colour affect the growth, survival and foraging behaviour
of Atlantic cod (Gadus morhua) larvae? Aquaculture 277(34):197-202.
Pea, R., S. Dumas, R. Saldivar-Luci, G. Garcia, A. Trasvia and D.
Hernndez-Ceballos. 2004. The effect of light intensity on first
feeding of the spotted sand bass Paralabrax maculatofasciatus
Steindachner larvae. Aquaculture Research 35:345-349.
Puvanendran, V. and J. Brown. 1998. Effect of light intensity on
the foraging and growth of Atlantic cod larvae: interpopulation
difference? Marine Ecology Progress Series 167:207-214.
Puvanendran, V. and J. Brown. 2002. Foraging, growth, and
survival of Atlantic cod larvae in different light intensities and
photoperiods. Aquaculture 214:131-151.
Rahmann H., G. Jeserich and I. Zeutzius. 1979. Ontogeny of
visual acuity of rainbow trout under normal conditions and light
deprivation. Behavior 68:315-322.
Raymond, P.A. 1985. Cytodifferentiation of photopreceptors
in larval goldfish: delayed maturation of rods. Journal of
Comparative Neurology 236:90-105.
Ruchin, A.B. 2004. Influence of colored light on growth rate of
juveniles of fish. Fish Physiology and Biochemistry 30:175-178.
Suthers, I.M. and S. Sundby. 1996. Role of the Midnight sun:
comparative growth of pelagic juvenile cod (Gadus morhua)
from the Arcto-Norwegian and a Nova Scotian stock. ICES
Journal Marine Science 53:827-836.
Tamazouzt, L., B. Chatain and P. Fontaine. 2000. Tank wall colour
and light level affect growth and survival of Euroasian perch
larvae (Perca fluviatilis L.). Aquaculture 182:85-90.
Trippel, E.A. and S.R.E. Neil. 2002. Effect of photoperiod and
light intensity on growth and activity of juvenile haddock
(Melanogrammus aeglefinus). Aquaculture 217:633-645.
Volsett, K.W., A. Folkvord and H.I. Browman. 2011. Foraging
behaviour of larval cod (Gadus morhua) at low light intensities.
Marine Biology 58:1125-1133.
Wilds, G.R. and M.I. Muoneke. 2001. Climate related and
morphoedaphic correlates of growth in white bass. Journal of
Fish Biology 58:453-461.
Yoon, H, J. Hwang and S. Choi. 2010. Effect of light intensity on
first feeding of the chub mackerelScomber japonicuslarvae.
Animal Cells and Systems 14:125-128.
Zeutsius, I. and H. Rahmann. 1984. Influence of dark-rearing on the
ontogenic development of Arotherodon mossambicus (Cichlidae,
Teleostei): I. Effects on body weight, body growth pattern,
swimming activity, and visual acuity. Experimental Biology
43:77-85.

W W W.WA S .O R G

W O R L D AQ UACU LT U R E

S E P T E M B E R 2 015

29