Partridge Et Al., 2016 - The Performance of Larval Seriola Lalandi Valenciennes 1833 is Affected by The

Aquaculture Research, 2017, 48, 1260–1268 doi:10.1111/are.
12967
The performance of larval Seriola lalandi

(Valenciennes, 1833) is affected by the taurine
content of the Artemia on which they are fed
Gavin J Partridge1,2 & Lindsey D Woolley1

1
Australian Centre for Applied Aquaculture Research, Challenger Institute of Technology, Fremantle, WA, Australia
2
Fish Health Unit and Freshwater Fish Group, School of Veterinary & Life Sciences, Murdoch University, Murdoch, WA,
Australia
Correspondence: G J Partridge, Australian Centre for Applied Aquaculture Research, Challenger Institute of Technology, 1 Fleet
Street, Fremantle, WA 6160, Australia. E-mail: gavin.partridge@challenger.wa.edu.au
yellowtail kingfish larvae may have a lower

Abstract
requirement and/or a reduced tolerance to excess
This study describes the effects of feeding taurine- dietary taurine than juveniles.
supplemented Artemia on the growth, survival,
whole body taurine content and jaw malformation Keywords: larviculture, dietary taurine, Artemia
rate of larval yellowtail kingfish Seriola lalandi. Lar- enrichment, marine finfish
vae were fed rotifers containing no supplemental
taurine from 3 to 15 day post hatch (dph) and
Introduction
Artemia co-enriched with taurine from 12 to 22
dph. Artemia were supplemented at concentrations Yellowtail kingfish Seriola lalandi is an established
of either 0, 0.8, 1.6, 2.4, 3.2 or 4.0 g of tau- aquaculture species in Japan and Australia
rine L1 during the 18 h HUFA enrichment pro- (Nakada 2002; Fielder 2013) and is being investi-
cess. Taurine content in the Artemia increased gated in many other countries and regions includ-
from 0.76 0.04% DW in those without supple- ing New Zealand (Poortenaar, Hooker & Sharp
mentation to 3.95 0.17% DW in those supple- 2001), the Americas (Benetti, Nakada, Shotton,
mented at 4.0 g L1. Survival rates of larval Poortenaar, Tracy & Hutchinson 2005) and Eur-
yellowtail kingfish were significantly lower in all ope (Abbink, Blanco Garcia, Roques, Partridge,
taurine-supplemented treatments compared to the Kloet & Schneider 2012). Unlike Japan, grow-out
unsupplemented control. Growth was significantly production in Australia is reliant upon hatchery
improved in those larvae fed taurine-supplemented produced juveniles and production is somewhat
Artemia; however, we cannot attribute this constrained by relatively low larval survival rates
improvement solely to taurine, as improved and juvenile quality, particularly jaw malforma-
growth may have been a function of the reduced tions (Cobcroft, Pankhurst, Poortenaar, Hickman
survival, and therefore increased prey availability, & Tait 2004). The rotifers and Artemia used to feed
in these treatments. The whole body taurine con- yellowtail kingfish larvae contain lower concentra-
tent of larvae fed unsupplemented Artemia was sig- tions of many nutrients and trace elements than
nificantly lower (1.85 0.03% DW) than those the wild zooplankton on which marine fish larvae
fed supplemented Artemia, which did not differ naturally feed, including taurine, and such defi-
from each other (pooled average 2.48 0.03% ciencies may be at least partially responsible for
DW), suggesting either a functional excretion such malformations (Cobcroft et al. 2004).
mechanism is in place or that this represents the Taurine is a neutral b-amino acid. It differs from
saturation value for larvae of this age. Jaw malfor- most amino acids in that it lacks a carboxyl group
mation rates were not affected by Artemia taurine and does not form peptide bonds, but it is the most
content. The results of this research suggest abundant free amino acid in animal tissues,
1260 © 2016 John Wiley & Sons Ltd

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Aquaculture Research, 2017, 48, 1260–1268 Artemia taurine content affects S. lalandi larvae G J Partridge & L D Woolley
including in marine fish larvae, accounting for up older fish due to the development of their organ
to 50% of the free amino acid pool (Conceicao, systems and based on the fact that egg and yolk-
van der Meeren, Verreth, Evjen, Houlihan & Fyhn sac larvae contain high levels of taurine (Pinto,
1997). It is found throughout the body in muscle, Figueira, Ribeiro, Yufera, Dinis & Arag~ ao 2010;
brain, ocular tissues, intestines, plasma and blood Pinto, Figueira, Santos, Barr, Helland, Dinis & Ara-
cells (Ripps & Shen 2012; El-Sayed 2013). It is g~ao 2013). These factors suggest that taurine sup-
involved in many different physiological processes plementation may be necessary for the larviculture
such as digestion via bile acid metabolism and the of yellowtail kingfish. Indeed, preliminary studies
regulation of blood cholesterol levels, neuromodu- with this species have demonstrated a benefit of
latory actions, cardiac Ca2+ modulation, osmoreg- feeding taurine-enriched rotifers (Rotman, Stuart &
ulation, vision, renal function, brain development Drawbridge 2012); however, studies have not
and reproduction (Ripps & Shen 2012). It acts as been conducted during the Artemia feeding stage.
a broad-spectrum cytoprotective agent and antiox- In a preliminary, unpublished study we found
idant (Ripps & Shen 2012) and has been found to reduced survival of yellowtail kingfish larvae fed
improve gut development (Li, Mai, Trushenski & Artemia enriched with taurine at the concentration
Wu 2009). It is considered a growth promoter in of 4 g L1 as per the methods of Salze et al.
many fish species possibly due to its role in (2011). The aim of this study was therefore to
enhancing the absorption of lipids and lipid-soluble confirm these findings and investigate the effect of
vitamins. Furthermore, and of potential impor- lower taurine enrichment concentrations on the
tance to Seriola larviculture, is taurine’s role in taurine content of Artemia and the subsequent
bone growth via the stimulation of bone-forming effect these Artemia have on the growth, survival,
osteoblasts and preventing formation of bone- jaw malformation rate and whole body taurine
degrading osteoclasts (Salze, Craig, Smith, Smith & content of yellowtail kingfish larvae.
McLean 2011).
Because taurine is not involved in protein syn-
Materials and methods
thesis, it is often considered to be ‘non-essential’,
however based on its many physiological roles and
Larval fish rearing system
broad distribution throughout the body, taurine is
clearly a critical nutrient (Ripps & Shen 2012). Fertilized yellowtail kingfish eggs were sourced
Taurine is certainly essential for those species from captive broodstock held at Clean Seas Tuna,
which lack the ability to produce it from its pre- Arno Bay (South Australia) and transported to the
cursors, L-cysteine and methionine, due to a lack Australian Centre for Applied Aquaculture
of cysteinesulfinic acid decarboxylase (CSD) activ- Research (Western Australia). The eggs were
ity (Ripps & Shen 2012). Many pelagic marine hatched in a 1000 L incubator at 22°C. After
species fall into this category and while the CSD hatching, 1 day post hatch (dph) larvae were ran-
activity of yellowtail kingfish has not been studied, domly stocked into twelve 300 L tanks at 60 lar-
Japanese yellowtail Seriola quinqueradiata (Tem- vae L1. All 12 tanks were treated equally during
minck & Schlegel) has been demonstrated to be the first 12 days. The rearing tanks were part of a
completely lacking in this enzyme (Yokoyama, flow-through system supplied with filtered seawa-
Takeuchi, Park & Nakazoe 2001). The rotifers and ter [34 g L1 with an exchange rate of 54 L h1
Artemia used to culture Seriola sp. contain lower (400% daily water exchange)] in each tank. Each
concentrations of taurine than the zooplankton on of four rearing tanks were floated in three 5000 L
which they would naturally feed. The taurine con- tanks to maintain the water temperature at 24°C
tent of wild zooplankton, for example, ranges from throughout the experiments. All tanks were com-
0.58% to 1.77% DW, whereas Artemia have been pletely independent of each other and no mixing
reported to contain only 0.63–0.83% DW (van der of water between them was possible. Two airstones
Meeren, Olsen, Hamre & Fyhn 2008; Yamamoto, were placed in each tank to maintain the dissolved
Teruya, Hara, Hokazono, Hashimoto, Suzuki, Iwa- oxygen levels close to saturation. During the first
shita, Matsunari, Furuita & Mushiake 2008) and 15 days a diffused metal halide light (400 W)
rotifers even less at 0.04–0.19% DW. Further- above each 5000 L tank provided a surface light
more, it has been hypothesized that larval fish intensity of 4600 1250 lux at the centre of
may have a higher requirement for taurine than each rearing tank for a photoperiod of 12 h light
© 2016 John Wiley & Sons Ltd, Aquaculture Research, 48, 1260–1268 1261
Artemia taurine content affects S. lalandi larvae G J Partridge & L D Woolley Aquaculture Research, 2017, 48, 1260–1268
(08:00 to 20:00 hours) and 12 h dark. Microalgal stereomicroscope one hour after the first feed on
paste (Nannochloropsis sp., Reed Mariculture, San 13, 15, 17 and 19 dph in squash-mounted fresh
Jose, CA, USA) was automatically dosed into the larvae. The trial was terminated on 23 dph and
rearing tanks during daylight hours to maintain larvae from each tank were hand counted to
turbidity within the range of 1.7 to 1.9 NTU determine the survival. One hundred larvae from
(equivalent to a Secchi disc depth of 55–60 cm) each tank were anesthetized without recovery,
from initial stocking until the end of the rotifer and fixed in 10% formaldehyde then transferred to
feeding stage. On 15 dph, microalgal additions 70% ethanol for jaw deformity assessment accord-
ceased and the light intensity was reduced to ing to methods described by Cobcroft et al. (2004).
800 150 lux at the centre of each rearing tank. Jaws classified as a commercial cull by the indus-
Between 3 and 12 dph, larvae were fed only try are represented as a percentage of the total
rotifers enriched with SPRESSO (INVE Aquacul- number of larvae assessed. The remaining larvae
ture, Dendermonde, Belgium) and without taurine from each tank were pooled, rinsed to remove sea-
enrichment under the hybrid feeding protocol water and frozen for analyses of taurine as
described by (Woolley & Partridge 2015). From 12 described above. Following the freeze-drying and
dph, feeding on taurine-enriched Artemia grinding of these pooled larvae a 2 g subsample
metanauplii began. Artemia were enriched with was analysed for taurine as described above.
SPRESSO (INVE Aquaculture) according to manu-
facturer’s directions. Artemia were co-enriched
Statistics
with taurine (Henan Aowei International, Zhengz-
hou, China) during the 18 h enrichment period at One-way ANOVA was used to determine differences
one of six concentrations 0, 0.8, 1.6, 2.4, 3.2 and between treatments in final growth of larvae, jaw
4.0 g TAU L1 following the method of Salze et al. deformity rates and larval taurine uptake. A
(2011). During 12 to 15 dph, larvae were co-fed repeated measures ANOVA was used to determine
rotifers and Artemia. Treatments were randomly the effect of larval age and Artemia enrichment
allocated across the 12 experimental tanks, with treatment on the number of Artemia consumed.
duplicate larval rearing tanks per treatment. Arte- Regression analysis was used to determine the
mia were fed to larvae according to the adaptive effect of dosage concentration on the taurine
feeding method described by Woolley, Partridge uptake in Artemia and the relationship between
and Qin (2012). taurine content in Artemia and larval taurine
whole body content. Data were arcsine trans-
formed where necessary to ensure homogeneity of
Sampling protocol
variance. Significance was set at P < 0.05 and val-
Sub-samples (ca. 100 g) of Artemia from each ues are presented as mean SE. All statistical
enrichment tank were taken on two different days, analyses were performed using IBM Statistics 20
rinsed in freshwater, frozen and then freeze-dried. (Release 20.0, Chicago, IL, USA).
Total taurine was analysed as part of total amino
acid profile on each sample via HPLC following
Results
homogenization then hydrolysis in 6 M HCl with
0.5% phenol for 24 h at 110°C according to Ray- The purity of the taurine was measured at
ner (1985) and Barkholt and Jensen (1989). Pur- 99.55% and all heavy metals were below their
ity of the taurine was also determined via this respective detectable limits.
method by comparing against a pure taurine stan- Artemia taurine content was significantly
dard (T-0625; Sigma Aldrich). Heavy metal analy- affected by enrichment concentration (P < 0.001;
sis of the taurine was conducted by preparing an Fig. 1). Those Artemia not receiving taurine sup-
acidified 1% solution in deionized water following plementation had a total taurine content of
analysis on an Agilent 730 Axial Simultaneous 0.76 0.04% DW, significantly lower than all
CCD ICP-OES. treatments receiving supplementation. Total Arte-
Larval dry weight was assessed on 23 dph on mia taurine content increased from 1.79 0.27%
20 randomly selected larvae per tank. Individual at the lowest concentration of 0.8 g L1 to 3.77
Artemia were quantified in the larval guts (5 per at 3.2 g L1. Artemia taurine plateaued at an
tank) by counting undigested Artemia eyes under a enrichment concentration of 3.2 g L1, with no
1262 © 2016 John Wiley & Sons Ltd, Aquaculture Research, 48, 1260–1268
Figure 2 Final survival (% mean SE, n = 2) of yel-

lowtail kingfish larvae at 23 days post hatch fed Arte-
Figure 1 Relationship between taurine enrichment mia co-enriched with increasing concentrations of
concentration (g L1) and Artemia taurine content (% taurine. Symbol asterisk indicates significant differences
dry weight) following an 18-h enrichment period between treatments.
(R2 = 0.99, Artemia taurine content (%
DW) = 0.76 + 4.35 9 (1 exp (0.34 9 taurine concentra-
tion rate)
).
significant difference in Artemia taurine content

between a concentration of 3.2 and 4.0 g L1
(3.77 0.13% and 3.95 0.16% DW respec-
tively). The relationship between the taurine con-
tent of the Artemia and the taurine enrichment
concentration was described by the following
equation with an R2 of 0.99.
Figure 3 The final dry weight (mean SE, n = 2) of
yellowtail kingfish larvae at 23 days post hatch fed at
Artemia taurine contentð%DWÞ ¼ 0:76 þ 4:35
ð1Þ varying levels of taurine co-enrichment during the
ð1 expð0:34taurineconcentrationÞ Þ Artemia feeding phase. Different superscripts indicate
significant difference between treatments.
Larval survival to 23 dph was significantly
average final dry weight of the larvae was strongly
affected by treatment (Fig. 2). Survival in the con-
and negatively correlated with their survival
trol (0 g L1) treatment (10.4 1.1%), was sig-
(R2 = 0.81; Fig. 4). Artemia ingestion by larvae
nificantly higher than all taurine-enriched
increased significantly with age (P > 0.001) but
treatments, which did not differ from each other
was equal across all treatments (P = 0.05; Fig. 5).
(pooled average 4.7 1%; P = 0.001). As a result
Larval whole body taurine content was signifi-
of the significant differences in survival, final lar-
cantly affected by Artemia taurine content
val density was also significantly affected by treat-
(P < 0.001). Those larvae receiving unsupple-
ment (P = 0.001). Larval density at the end of the
mented Artemia had a significantly lower whole-
trial in the control treatment (5.5 0.4 lar-
body taurine content (1.85 0.03% DW) than all
vae L1) was more than double that in all tau-
taurine-supplemented treatments, which did not
rine-enriched treatments, which did not differ from
differ from each other (pooled average
each other (pooled average 2.5 0.18 lar-
2.48 0.03% DW). The relationship between the
vae L1).
taurine content of the Artemia and the whole body
Final larval dry weights were significantly
taurine content of the larvae was described by the
affected by enrichment concentration (P = 0.01
following equation with an R2 of 0.98 (Fig. 6).
and P = 0.04 respectively). Those larvae fed Arte-
mia without taurine enrichment were significantly
smaller (3.8 0.3 mg DW; Fig. 3) than those Larval whole body taurine ¼ 0:79 þ 1:75
ð2Þ
receiving Artemia at all enrichment concentra- ð1 expð1:2ArtemiataurinecontentÞ Þ
tions, except those enriched at 2.4 g L1. The
we selected were based on the study by Salze et al.

(2011) in which larval cobia Rachycentron cana-
dum (Linnaeus) were fed rotifers and Artemia
which had both been enriched with taurine at a
concentration of 4.0 g L1. While the taurine con-
tent of the live foods were not presented by these
authors, they were reported in a later publication
as being ca. 0.08% and 0.23%, on a wet weight
basis for unsupplemented and supplemented Arte-
mia respectively (Salze, McLean & Craig 2012). To
convert these values to % DW, we measured the
water content of Artemia at 90% and subsequently
calculated these values to equate to ca. 0.8% DW
Figure 4 Relationship between the final dry weight and 2.3% DW respectively. While this level in the
(mg larvae1) and survival of yellowtail kingfish larvae unsupplemented Artemia is equivalent to that
(R2 = 0.81). reported here, the level we achieved in Artemia
supplemented at 4.0 g L1 was much higher than
There was no effect of taurine enrichment concen-
achieved by Salze et al. (2012) (3.95% DW cf.
tration on jaw deformity levels (P = 0.77; Fig. 7).
2.3% DW). These differences may have been due
Deformities considered a ‘commercial cull’ by the
to differences in enrichment time, as it has been
industry were 17.3 1.5% (pooled mean SE).
demonstrated that taurine uptake by live feeds is
time-dependent (Chen, Takeuchi, Takahashi,
Discussion
Tomoda, Koiso & Kuwada 2004, 2005). Further
This appears to be the first study investigating the studies are therefore required to fully elucidate the
response of live foods to a range of different interactive effects of enrichment time and concen-
taurine enrichment concentrations. Our results tration on the taurine content of live foods.
demonstrate that Artemia effectively take up tau- All larvae receiving taurine-enriched Artemia in
rine during the enrichment process in a relation- the current trial experienced significantly lower
ship that is linear at low concentrations then survival than those receiving unsupplemented
plateaus at the highest concentrations tested. The Artemia and this could not be attributed to impuri-
Artemia taurine levels we achieved in all but the ties in the taurine used. This finding is consistent
lowest enrichment concentration were higher than with our preliminary unpublished study where
typically seen in wild zooplankton (range 0.58– Artemia were enriched at 4.0 g L1. The majority
1.70% DW; van der Meeren et al. 2008; Yama- of published studies dealing with the taurine
moto et al. 2008). The rates of supplementation enrichment of live foods for marine fish larvae
Figure 5 Artemia ingestion, mea-

sured as the number of undigested
Artemia eyes per larval gut 1 h
after the first feed, of yellowtail
kingfish larvae from 13 to 19 days
post hatch. Five larvae per sample
time per replicate tank, values are
mean SE (n = 2).
1.41% (Pinto et al. 2013). On the other hand,

Salze et al. (2011) reported that cobia larvae fed
both taurine-enriched rotifers (estimated taurine
content 0.5% DW) and Artemia (estimated taurine
content 2.3% DW) experienced ca. four times
greater survival than those fed unsupplemented
live feeds. These authors did not separate the per-
formance of the cobia larvae between the rotifer
and Artemia feeding phases and the reasons for
the very different response of cobia larvae to sup-
plemental taurine compared with both the yellow-
tail kingfish in the current trial and the other
aforementioned studies on marine fish larvae is
Figure 6 Correlation between the taurine content (% dry unclear.
weight) in Artemia and yellowtail kingfish larvae fed Arte- Despite a lack of published studies showing a
mia co-enriched at increasing concentrations of taurine negative impact of taurine on the survival of mar-
(R = 0.98, Larval whole body taurine =
ine fish larvae, there does appear to be an emerg-
0.79 + 1.75 9 (1 exp(1.2 9 Artemia taurine content)).
ing body of evidence that live feed taurine
contents higher than those reported above in the
studies on Japanese flounder, red sea bream, Paci-
fic cod and gilthead sea bream can be detrimental
to marine fish larvae. In two independent trials
(performed in successive years), Koven, Nixon,
Azouli, Allon, Gaon, El Sadin, Falcon, Besseau,
Escande and Tandler (2014) fed rotifers with tau-
rine contents of 0.11% DW (unenriched), 0.44%
DW or 0.64% DW to Atlantic bluefin tuna Thun-
nus thynnus (Temminck & Schlegel) larvae. In both
studies those larvae fed the highest level of taurine
exhibited lower survival than those fed the inter-
mediate concentration of taurine. Similarly, Hawk-
Figure 7 Jaw deformity levels in yellowtail kingfish yard, Laurel, Barr et al. (2014) presented data
fed varying amounts of taurine-enriched Artemia from
showing a reduction in the survival of yellowtail
12 to 23 days post hatch. Levels represent deformities
kingfish larvae when fed rotifers enriched to con-
that would represent a commercial cull by industry.
tain 1.5% DW of taurine relative to the control of
Values are mean SE (n = 2).
unsupplemented rotifers. The lowest enrichment
concentration tested in the current study
report no benefit or negative impact on survival. (0.8 g L1) yielded Artemia with a taurine content
For example, survival of Japanese flounder Par- of 1.8% DW; higher than those which caused a
alichthys olivaceus (Temminck & Schlegel), red sea negative impact on Atlantic Bluefin and yellowtail
bream Pagrus major L., Pacific cod Gadus macro- kingfish larvae by the former two authors.
cephalus L. and northern rock sole Lepidopsetta In terms of larval growth, most published studies
polyxystra (Orr & Matarese) larvae were not report a positive benefit of enriching live feeds with
improved by increasing the taurine content of roti- taurine. For example in the aforementioned trials
fers to between ca. 0.3% and 0.45% DW (Chen on red sea bream, Japanese flounder, Pacific cod
et al. 2004, 2005; Matsunari, Arai, Koiso, and northern rock sole, all species grew significantly
Kuwada, Takashi & Takeuchi 2005; Hawkyard, faster when fed rotifers enriched to contain between
Laurel, Barr, Stuart, Drawbridge & Langdon 0.3% and 0.45% DW of taurine than those larvae
2014). Likewise there was no benefit to the sur- fed unsupplemented rotifers (Chen et al. 2004,
vival of gilthead sea bream Sparus aurata L. larvae 2005; Matsunari, Arai et al. 2005; Hawkyard, Lau-
when fed rotifers whose taurine content had been rel & Langdon 2014). Senagalese sole Solea sene-
increased from 0.88% of total amino acids to galensis (Kaup) larvae fed microcapsules containing
taurine grew faster than those fed microcapsules Chen, pers. comm., 2014). While supplementation
without taurine (Pinto et al. 2010) and cobia larvae with taurine may have been responsible for the
grown on taurine-enriched live foods grew signifi- improved growth of larvae in those treatments,
cantly faster than those receiving unsupplemented further studies in which reduced survival does not
live foods (Salze et al. 2011). lead to significant differences in prey availability
While we also achieved significantly greater lar- and final larval density will be required to prove
val growth in all taurine-supplemented treatments, this hypothesis.
we are unable to attribute this difference solely to Given that taurine plays a role in bone growth
taurine, as such differences may also be at least and development (Salze et al. 2011), we hypothe-
partially attributable to differences in prey avail- sized that taurine supplementation may reduce the
ability and/or larval density as a result of the sig- incidence of jaw malformations in yellowtail king-
nificantly different survival rates between fish larvae. That no significant differences in jaw
treatments. On the basis of previous data we sug- malformations were found demonstrates there to
gest that larvae should not have been food limited. be no beneficial effects of the taurine supplementa-
Based on the survival rates achieved in each treat- tion regimes employed in this trial. Further studies
ment and with the feeding regime employed, we investigating the effects of lower Artemia taurine
have calculated that larvae in the taurine-enriched supplementation concentrations may yield different
treatments would have had access to 416 30 results.
Artemia per larvae per meal at the end of the trial, Despite significant differences in the taurine con-
while those in the control treatment had access to tent of Artemia at all but the two highest taurine
182 13 Artemia per larvae per meal. While this supplementation rates, there were no significant
difference is significant, those larvae in the control differences in the whole body taurine contents of
treatment had access to a similar number of Arte- the larvae that consumed these different supple-
mia to those in Woolley et al. (2012), which mented Artemia. This suggests that either larvae of
received approximately 140 Artemia per larvae per this age already possess the functional mecha-
meal at the same age. That there was no correla- nisms required for excreting taurine in excess of
tion between survival and larval size in this afore- their requirements, or that their bodies are fully or
mentioned trial, even with a lower feed rate, super-saturated with taurine. The former hypothe-
suggests that larvae in the current trial should not sis would imply that the bodies of these larvae
have been food limited. Furthermore, the lack of have reached a state of homoeostasis and that any
difference in the measured Artemia ingestion rates taurine beyond the larvae’s requirements is being
in the current trial also supports our hypothesis excreted; a process that may have been demand-
that larvae were not food limited. We therefore ing on these larvae and reduced their survival.
consider that differences in larval density may The negative impact on survival in all supple-
have played a more important role than food mented treatments also supports our alternative
availability in any differences in fish size not attri- hypothesis that the bodies of larvae in all supple-
butable to taurine. Larval density at the end of the mented treatments are fully or super-saturated
trial in the control treatment (5.5 0.4 lar- and subsequently that these whole body levels are
vae L1) was more than double that in all tau- in excess of the larvae’s requirements. A compar-
rine-enriched treatments (pooled average ison of the whole body taurine contents of wild
2.5 0.18 larvae L1) and the average final dry juvenile conspecifics supports this theory. Larvae
weight of the larvae was strongly correlated with fed taurine-supplemented Artemia in this study
their final larval density (R2 = 0.81). Density- had a higher whole body taurine content (2.48%
dependent growth has been described in other DW) than for both 30 mm wild juvenile Japanese
marine fish larvae. Increasing the density of yel- yellowtail (2.3% DW) (Matsunari, Takeuchi, Taka-
lowfin tuna Thunnus albacares (Temminck & Sch- hashi & Mushiake 2005) and for wild juvenile
legel) larvae from 2 to 18 L1, for example, amberjack Seriola dumerili (Risso) (2.1% DW) of
resulted in growth reductions of up to 35% (Mar- 28–44 mm in length (Yamamoto et al. 2008).
gulies, Scholey, Wexler, Olson, Suter & Hunt Given that taurine is involved in many different
2007). Density-dependent larval growth was also physiological mechanisms, it is conceivable that
observed in unpublished trials investigating the one or more of these mechanisms are being over-
effects of larval density in yellowtail kingfish (B. whelmed by excessive taurine in underdeveloped
larvae. Larvae such as those used in the current 754). The authors thank Clean Seas Tuna for pro-
trial may therefore have a lower requirement and/ viding the eggs. The authors also thank the Aus-
or a reduced tolerance to excess taurine than juve- tralian Centre for Applied Aquaculture Research
nile fish. For example, juvenile Japanese flounder for use of facilities and support through the larval
fed zooplankton (mysids) have a similar whole- rearing.
body taurine content (2.4% DW) to the aforemen-
tioned wild Japanese yellowtail and amberjack
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Aquaculture Research, 2017, 48, 1260–1268 doi:10.1111/are.

The performance of larval Seriola lalandi

Gavin J Partridge1,2 & Lindsey D Woolley1

yellowtail kingfish larvae may have a lower

1260 © 2016 John Wiley & Sons Ltd

Figure 2 Final survival (% mean SE, n = 2) of yel-

significant difference in Artemia taurine content

we selected were based on the study by Salze et al.

Figure 5 Artemia ingestion, mea-

1.41% (Pinto et al. 2013). On the other hand,

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