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Aquaculture Nutrition 2003 9; 231^237

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Development of a pelleted feed for juvenile tropical spiny


lobster (Panulirus ornatus): response to dietary protein
and lipid
D.M. SMITH, K.C. WILLIAMS, S. IRVIN, M. BARCLAY & S. TABRETT
CSIRO Marine Research, Cleveland Qld., Australia

Introduction
Abstract
Of all of the rock or spiny lobster species, the tropical spiny
Critical to the development of a cost-effective feed for the
lobster, Panulirus ornatus is probably the best candidate for
tropical spiny lobster Panulirus ornatus is knowledge of its
aquaculture as it has the shortest oceanic larval development
response to the protein and lipid (or energy) content of the
phase (4–6 months) and the fastest growth rate (1 kg within
feed. An experiment of 12 weeks duration was carried out to
2 years after hatching) (Phillips et al. 1992; Pitcher et al.
examine growth responses of juvenile lobsters to pelleted
1995; Dennis et al. 1997; Butler & Herrnkind 2000). Where
diets that provided six crude protein (CP) levels [320–
P. ornatus postpueruli and early juveniles (or ÔseedÕ) are
600 g kg)1 dry matter (DM)] and two lipid levels (nominally
legally collected from the wild, lobster culture has become a
60 and 100 g kg)1 DM). Lobsters (mean initial weight of
profitable and flourishing aquaculture industry, with further
1.8 g) were held in groups of nine or 10 animals in
expansion being limited only by the availability of wild seed
24 · 350 L tanks, fed twice daily at a restricted level, and
(Srikrishnadhas & Rahman 1995; Lovatelli 1997; Tuan et al.
maintained at 28 C. Maximal growth responses occurred at
2000). In Australia, where collection of wild lobster seed for
dietary CP contents of 474 g kg)1 for the 60 g kg)1 lipid
commercial on-growing is not generally permitted, estab-
series and 533 g kg)1 for the 100 g kg)1 lipid series. A second
lishment of P. ornatus aquaculture may depend on com-
experiment, of 4 weeks duration, compared two dietary
mercially viable hatchery technology being developed for
treatments: a mixture of two of the best diets from the first
large-scale propagation of the species. While culture of
experiment, and a commercial shrimp (Penaeus japonicus)
P. ornatus from egg through the larval phase to puerulus has
feed. Lobsters were held under the same experimental con-
not been reported, this is an area of active research in Aus-
ditions as in the first experiment, but were fed to excess twice
tralia and initial results are very encouraging (C. Jones, pers.
daily. Their growth was significantly greater (P < 0.05) on
comm.). Moreover, the recent success in propagating other
the shrimp feed (0.68 g week)1) than on the laboratory-
slower-developing species of spiny lobsters within the genera
pelleted diets used in the main study (0.32 g week)1). The
Panulirus, Palinurus and Jasus (Illingworth et al. 1997;
results indicate that the optimal dietary protein and lipid
Kittaka 1997a,b; Matsuda & Yamakawa 2000) augur well
content of the diet for P. ornatus is about 530 and 100 g kg)1,
for successful propagation of P. ornatus.
respectively.
One of the obstacles to sustainable spiny lobster aqua-
culture is the lack of a suitable formulated feed that will
KEY WORDS: feeding, nutrition, protein : energy, rock
enable juveniles to be reared economically to a marketable
lobster
size. Natural food items such as fresh fish, crustaceans and
molluscs have proved to be suitable for culture of juvenile
Received 15 May 2002, accepted 7 October 2002
spiny lobsters under laboratory (Dennis et al. 1997; Hooker
Correspondence: D.M. Smith, CSIRO Marine Research, PO Box 120,
et al. 1997; James & Tong 1997) and commercial (Tuan et al.
Cleveland Qld. 4163, Australia. E-mail: [email protected]
2000) conditions in Vietnam. However, lobster culture based
on feeding fresh natural foods is unlikely to be permitted by
resource managers in Australia because of feed wastage and

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 2003 Blackwell Publishing Ltd 231


232 D.M. Smith et al.

the resulting release of nutrients to the environment. Food Table 1 Formulation of the terminal diets1 of each series of diets
conversion ratios for lobsters in Vietnam on fresh natural used in the examination of protein and protein : energy responses of
juvenile Panulirus ornatus in Experiment 1. Diet label denoting lipid
food have been reported to be as high as 28 : 1 (Tuan et al.
(L) and protein (P) content on an Ôas fedÕ basis
2000). Moreover, collection of natural foods in the quantities
needed for large-scale commercial culture of lobsters would 60 g kg)1 lipid series 100 g kg)1 lipid series

be logistically difficult and prohibitively expensive in Ingredient 60L : 300P 60L : 550P 100L : 300P 100L : 550P
Australia. A recent economic analysis of the feasibility of Fishmeal (defatted) 2
133 494 133 494
J. edwardsii aquaculture in New Zealand showed it to be Fishmeal (68% CP) 287 284 288 284
unprofitable unless infrastructure and operating costs could Starch 339 18 333 10
Fish oil 11 6 53 46
be substantially reduced (Jeffs & Hooker 2000). Developing a
Mussel (dry equiv.)3 10 10 10 10
cost-effective formulated feed as an alternative to the feeding Crustacean meal 51 51 51 51
of fresh mussels was identified in their analysis as a priority, Diatomaceous earth 69 37 32 5
if the economic outlook of culturing spiny lobsters in New Other ingredients4 100 100 100 100

Zealand was to be improved. 1


Each lipid series consisted of six diets serially incremented between the
Initial feeding studies with P. ornatus, J. edwardsii and amounts in terminal formulations.
2
Fishmeal (68% CP) that was defatted using hexane/ethanol (2 : 1)
P. cygnus showed that each species would readily eat pelleted
extraction.
dry diets, but that they had marked preferences for certain 3
DM equivalent of incorporated fresh wet mussel and based on the
ingredients (Williams et al. 2000). The next step is to develop analysed DM content of 200 g kg)1.
inexpensive diets that are not only attractive to the animals
4
Comprised (g kg)1 of diet): cholesterol, 2; lecithin,12; carophyll pink (8%
astaxanthin, Roche), 0.9; choline chloride, 0.2; vitamin premix (as
but also satisfy their nutritional needs for rapid growth. recommended by Conklin, 1997), 2; Ascorbyl-2-polyphosphate (Roche,
Critical to this objective is determining the optimal dietary Stay-C), 1; Ethoxyquin (Banox E), 0.2; alginate (Manucol), 61; tri-Na-
protein and lipid specification for each species. Companion polyphosphate, 20; and chromic oxide, 0.5.

studies, using the same diets as used in this study on


P. ornatus, have been carried out with P. cygnus (Glencross Table 2 Chemical composition of key ingredients used in diet for-
et al. 2001) and J. edwardsii (Ward et al. 2002) to define the mulations for the protein/lipid response study with juvenile P. ornatus
in experiment 1
spiny lobster’s responsiveness to dietary changes in protein
and lipid concentration. Fishmeal1 Fishmeal2 Crustaean
(defatted) (68% CP) Mussel3 meal4

g kg)1 as used
Methods Dry matter 889 909 100 920
g kg)1 DM basis
Experiments Ash 207 189 142 329
Crude protein 776 725 490 401
Two lobster-feeding experiments were carried out. The first Total lipid 16 116 100 74
experiment (experiment 1), of 12 weeks duration, examined 1
Peruvian fishmeal, 68% CP, extracted with hexane and ethanol (2 : 1) as
responses of juvenile P. ornatus to diets that provided six described.
2
protein levels at each of two lipid levels in a factorial design. Peruvian fishmeal, 68% CP., Ridley AgriProducts, Narangbar,
Queensland, Australia.
For experiment 2, two of the best diets from experiment 1 3
New Zealand GreenshellTM mussels, Sealord P/L, Nelson, New Zealand.
were combined (50 : 50) and compared over 4 weeks with an 4
Langostine meal,Tharos S.A., Chile.
extruded shrimp feed (Lucky Star, Taiwan Hung Kuo
Industrial Co., I-Lan, Taiwan), formulated for the kuruma and diatomaceous earth (to balance ash). Lipid concentration
shrimp, Penaeus japonicus. was manipulated by varying the inclusion of cod liver oil
(Melrose Laboratories, Box Hill, Victoria, Australia) at the
expense of starch. Fresh greenlip mussel Perna canaliculatus
Diet formulations
(New Zealand GreenshellTM mussel; Sealord P/L, Nelson,
Twelve diets were made up to provide six protein concentra- New Zealand) was included in all diets to enhance the
tions at either of two lipid concentrations (Tables 1 and 2). The attractiveness of the diet. Vitamin premix was supplied by
protein concentration was manipulated by varying the Rhone-Poulenc, Queensland, and ascorbyl-2-polyphosphate
inclusion of a de-fatted Peruvian fishmeal at the expense of (Stay-C; Roche, Sydney, Australia) by Ridley Agriproducts,
starch (B011C starch, Earlee Products, Queensland, Australia) Narangba Queensland. Carophyll Pink (8% astaxanthin) was

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P. ornatus response to dietary protein and lipid 233

donated by Roche, Australia. Other feed additives were polycarbonate sheeting covers. The tanks were supplied
obtained from Sigma-Aldrich, Sydney, Australia. aerated flowing seawater (500 mL min)1) at 28 C and
The defatted Peruvian fishmeal was prepared by three salinity between 33 and 35 g L)1.
successive extractions with a mixture of hexane and ethanol After acclimatization for 14 days to the experimental
(2 : 1) and then dried in a forced-draught fume hood at room conditions, lobsters were individually weighed (initial weight)
temperature until no odour of hexane could be detected. It and re-weighed at 4-weekly intervals until the experiment
was then reground in a hammer mill (Mikro-Pulverizer, NJ, ended at 12 weeks. They were fed their allocated diets (Table
USA) and sieved to pass through a 710 lm screen. 3) twice daily at 95% of satiety for the least preferred diet,
Diets were prepared by mixing the dry ingredients, after which had been determined during the acclimatization per-
which the homogenized fresh mussel, oil and sufficient water iod. This feeding rate (as a percentage of lobster biomass in
were added to form a soft dough [approximately 400 g kg)1 each tank) was maintained for the duration of the experi-
dry matter (DM)]. The dough was thoroughly mixed and ment, being adjusted after each 4-weekly weighing. About a
pressed through a 3 mm die attached to an electrohydraulic quarter of the daily ration was given in the morning (09:00
sausage filler (Hall Food Equipment, New South Wales, hours) and the balance in the afternoon (16:00 hours).
Australia) into a setting bath of 10% CaCl2 for 5 min. Mortalities were recorded daily and stocking density main-
The spaghetti-like strands were then dried overnight tained by introducing a tagged lobster of similar size; only
(850 g kg)1 DM) in a forced-draught fume hood and data from non-tagged animals were used in the ANOVA.
stored at )15 C until used. The pellets were subsequently At the end of experiment 1, 60 lobsters that had been fed the
reduced to approximately 5 mm in length. This appeared to three highest-protein diets were paired by weight and previous
be an appropriate size for the animals being fed, as the lob- diet treatment, and distributed to six pairs of tanks with five
sters were observed to ingest them with minimal fragmenta- lobsters per tank, so that lobsters in each pair of tanks were
tion and wastage. closely matched by weight. They were acclimatized to the
tanks for 2 weeks and fed a mixture of fresh mussel, the two
best-performing laboratory-pelleted diets (100L : 500P,
Experimental design, animals and management
100L : 550P) and kuruma shrimp pellets. Following this
Experiment 1 was a two-factor design with six dietary protein acclimatization, the lobsters were individually weighed (mean
levels and two lipid levels. The experiment was arranged as a of 3.9 ± 1.25 g) and one tank of each pair of tanks was ran-
balanced block design with each treatment randomly allo- domly assigned to one of the two dietary treatments for
cated to one tank within each of two blocks. Approximately
400 juvenile P. ornatus of 1–4 g were collected from Trinity Table 3 Chemical composition of the diets fed to juvenile P. ornatus
Inlet, Cairns, North Queensland, (1655¢S, 14545¢E) by in protein/lipid response study, experiments 1 and 2
divers and shipped to CSIRO Cleveland. The lobsters were Chemical composition
placed in tanks provided with aerated, flow-through filtered
DM Ash Protein Lipid Gross energy1
(20 lm) seawater at 28 C and salinity between 33 and
35 g L)1. For 8 weeks prior to the start of experiment 1, they Diet label (g kg)1 as fed) (g kg)1 DM) (MJ kg)1 DM)

were fed twice daily to satiety plus a slight excess on fresh 60L : 300P 792 236 323 62 15.64
greenlip mussel and a high-protein [640 g kg)1 crude protein 60L : 350P 791 262 403 66 15.89
60L : 400P 838 237 452 61 16.14
(CP), DM basis], kuruma shrimp feed.
60L : 450P 828 255 498 60 16.39
For experiment 1, the lobsters were sorted by weight into 60L : 500P 823 259 542 58 16.63
two groups from which 120 individuals of mean (±SD) 60L : 550P 807 269 611 58 16.88
weight of 1.2±0.27 g (block 1) and 108 individuals of 100L : 300P 861 190 330 107 17.34
100L : 350P 859 200 382 106 17.59
2.5±0.82 g (block 2), were randomly and equally assigned to 100L : 400P 855 208 435 106 17.84
12 experimental tanks per block, at stocking densities of 10 100L : 450P 849 214 488 108 18.09
and nine animals per tank, respectively. The experimental 100L : 500P 848 225 543 106 18.33
100L : 550P 846 235 598 105 18.58
tanks (1500 · 600 · 500 mm deep, 350 L capacity) were Shrimp diet2 945 162 640 122 20.20
constructed of fiberglass with a black gel coat on the inside
1
Derived by linear interpolation between values for extremity diets from
and the bottom treated to provide a non-skid surface; suffi-
each series.
cient clay bricks were placed in each tank to provide shelters 2
Commercial extruded kuruma shrimp diet that was fed in experiment 2
for the lobsters and the tanks were fitted with twin-walled only. DM ¼ dry matter.

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234 D.M. Smith et al.

experiment 2: (a) an equal mixture by weight of diets 100L : 500P Table 4 Productivity response means of juvenile P. ornatus in
and 100L : 550P (Tables 1 and 2); or (b) kuruma shrimp feed. Experiment 1
The lobsters were fed their respective diets twice daily to slight Productivity response
excess for 4 weeks after which they were re-weighed.
Initial Final Growth rate SGR Survival
Diet label1 weight (g) weight (g) (g week)1) (% day)1) (%)

Chemical and statistical analyses 60L : 300P 1.9 2.9 0.08 0.5 53
60L : 350P 1.9 3.6 0.14 0.8 58
Samples of finely ground raw ingredients or diets were ana- 60L : 400P 2.0 4.2 0.18 0.9 63
lysed in duplicate by standard laboratory methods essentially 60L : 450P 1.7 3.4 0.14 0.9 63
60L : 500P 1.8 3.7 0.16 0.8 53
in accordance with AOAC International (1999) recommen- 60L : 550P 1.8 2.8 0.09 0.6 69
dations, at either the CSIRO Marine Laboratory, Cleveland, 100L : 300P 1.9 2.7 0.07 0.5 79
or at the Animal Research Institute, Yerrongpilly (Tables 2 100L : 350P 1.7 3.6 0.15 0.9 44
100L : 400P 1.8 3.9 0.17 1.0 32
and 3). DM was determined by oven-drying at 105 C to 100L : 450P 1.8 3.5 0.14 0.8 74
constant weight; ash by ignition at 600 C for 2 h; total 100L : 500P 1.7 4.1 0.20 1.1 52
nitrogen (total N) by a macro-Kjeldahl technique on a Kjel 100L : 550P 1.9 4.4 0.21 1.0 69
?SEM 0.07 0.37 0.032 0.13 10.7
Foss automatic analyser using mercury in the digestion; CP
1
was calculated by multiplying total N by 6.25 irrespective of Refer toTable 1 for details.
the nature of the N. Total lipid (TL) was determined gravi- SGR ¼ specific growth rate.

metrically after a Bligh & Dyer (1959) extraction as modified


by Christie (1982). Gross energy (GE) was determined by lobsters improved curvilinearly with increased protein in the
isothermal bomb calorimeter using a microprocessor- diet. The protein content providing the maximum growth
controlled Leco AC 200 automatic bomb calorimeter (Leco rate varied according to dietary lipid content (Fig. 1). For the
Corp., St Joseph, MI, USA). The fatty acid compositions of 60 g kg)1 lipid series, the relationship was significant
the TL extracts of the diets used in experiment 2 were (P < 0.01) with the maximum response occurring at a dietary
determined as methyl ester derivatives (FAME). Lipids were CP content of 474 g kg)1 DM; for the 100 g kg)1 lipid series,
esterified by the method of Morrison & Smith (1964) and the relationship approached significance (P ¼ 0.10), with the
analysed by gas chromatography (GC) using a Hewlett maximum response occurring at a dietary CP content of
Packard 5890 capillary GC (Agilent Technologies, Mel- 533 g kg)1 DM.
bourne, Australia) with direct on-column injection and flame Despite the apparent difference between the 60 and
ionization detection. The FAME were separated on a 50 m 100 g kg)1 lipid series in the response to dietary protein
polar column (BP20, 0.33 mm i.d., 0.5 lm film thickness, (Fig. 1, Table 5), homogeneity tests of the slopes and inter-
Scientific Glass Engineering, Melbourne, Australia) with a cepts showed no significant difference (P > 0.05) between the
hydrogen carrier gas flowing at 2.7 mL min)1. two lipid series. The equation for the pooled data set showed
Data from experiment 1 were analysed as a two-way SGR significantly increased curvilinearly (P < 0.05) with
ANOVA (Snedecor & Cochran 1967) with statistical packages

(Queensland Department of Primary Industries, Queensland)


for balanced data (BALF) and regression (REGN), taking
into consideration the blocked design of the experiment.
Data was analysed further using the weight of the lobsters as
a covariate. For experiment 2, the data were analysed as a
one-way ANOVA. Percentage survival data were analysed as
the natural and arcsine-transformed values, but as the
transformation did not materially alter the significance of the
F-statistic, only the natural values are reported.

Results
Figure 1 Effect of dietary CP concentration on the SGR response
In experiment 1, survival rate was low but unaffected by the of spiny lobsters fed diets containing either 60 (m) or 100 (j) g kg)1
diet fed (Table 4). The specific growth rate (SGR) of the lipid for 12 weeks in Experiment 1.

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P. ornatus response to dietary protein and lipid 235

Table 5 Regression statistics describing the relationship between However, the weight gain was low, even on the best diet
SGR; Y of spiny lobsters and dietary CP (g kg)1 DM; X) for diets (100L : 550P).
containing either 60 or 100 g kg)1 dietary lipid in experiment 1
Lobsters fed the kuruma shrimp diet grew at twice the rate
Quadratic equation of the form: Y ¼ a + bX + cX2 and recorded a significantly higher survival than those fed the
Lipid
series (L) a b ?SEb c ?SEc R2 P best of the laboratory-pelleted diets (P < 0.05) (Table 6). The
gross nutritional composition of the two diets was sufficiently
60 L )2.589 0.0147 0.001663 )0.0000155 0.0000018 0.96 <0.01
100 L )1.988 0.0113 0.008464 )0.0000106 0.0000091 0.65 >0.05 similar (Table 3) not to account for the observed productivity
Pooled )2.437 0.0137 0.004609 )0.0000138 0.0000049 0.56 <0.05 differences. Although the fatty acid profiles of the two dietary
treatments in experiment 2 were broadly similar, the labor-
atory-pelleted diet had a higher content of highly unsaturated
Table 6 Productivity response means of juvenile P. ornatus in n-3 fatty acids, which one might speculate as being nutri-
experiment 2 tionally more favourable. Because of the difficulty in meas-
Productivity response uring feed consumption accurately, there are no reliable data
to indicate whether the productivity differences between the
Initial End Growth rate SGR Survival
Diet label weight (g) weight (g) (g week)1) (% day)1) (%) two diets were due to nutritional quality or feed intake dif-
ferences. However, as the survival rate of lobsters fed the
100L : 500/550P 3.8a 5.1b 0.32b 1.0b 80b
Shrimp diet1 3.9a 6.6a 0.68a 2.0a 100a kuruma shrimp diet was significantly higher than of those fed
?SEM 0.17 0.38 0.061 0.14 3.7 the laboratory-pelleted diet (100% vs. 80% respectively,
a,b
Within productivity traits, means without a common superscript letter Table 6), the latter diet was possibly nutritionally inadequate.
differ (P < 0.05). Alternatively, the laboratory-pelleted diet may have been lack-
1
Commercial extruded kuruma shrimp diet. ing in attractants, or lost its attractiveness far more rapidly
SGR ¼ specific growth rate.
than the kuruma shrimp diet, so that the lobsters were canni-
balizing other lobsters that were in a vulnerable condition,
increasing dietary CP with the maximum response occurring such as immediately after molting. Cannibalism of recently
at 495 g kg)1 DM CP (Fig. 1, Table 5). molted lobsters was observed to be the main cause of mortality
In experiment 2, lobsters fed the kuruma shrimp diet had a of juvenile P. ornatus in the study by Jones et al. (2001).
significantly higher survival rate and gained weight at twice The SGR of spiny lobsters fed the kuruma shrimp diet in
the rate of the lobsters fed the laboratory-pelleted diet (Table experiment 2 was higher (2.0% day)1) than that reported by
6). The fatty acid profiles of the combined 100L : 500P/ Jones et al. (2001) (1.56% day)1) for the same species held in
100L : 550P diet and the kuruma shrimp diet were broadly tanks and fed a mixture of a shrimp feed (for Penaeus
similar: saturated fatty acids comprising 33% (laboratory- japonicus, 580 g kg)1 CP) supplemented with shrimp flesh.
pelleted diet) and 36% (kuruma shrimp diet) of total fatty However, though the initial weight of lobsters in the two
acids; mono-unsaturated fatty acids comprising 15 and 35%; studies was similar (3.6 g vs. 3.3 g), in the latter study the
linoleic acid comprising 13 and 9%; linolenic acid 2% in both lobsters were on-grown to a much larger size. Phillips et al.
diets; eicosapentanoic acid 12 and 9%; and docosahexanoic (1992) reported that growth of postpuerulus P. ornatus held
acids comprising 16 and 10% of total fatty acids. under aquarium conditions and fed five times weekly
(presumably on a diet of natural foods) exhibited at an early
age, a sigmoidal increase in carapace length (CL), but
Discussion
thereafter carapace growth curves were well modelled by von
Lobster growth exhibited a clear dose response to dietary Bertalanffy relationships. Their data for laboratory-held
protein concentration (Fig. 1) with the benefit of the higher post-settlement juveniles show a CL increase from 7 mm at
lipid content being apparent in the high-protein diets. Max- placement to about 15 mm after 120 days and to 30 mm
imal growth responses occurred at dietary CP contents of after about 250 days. Using morphometric relationships for
474 g kg)1 DM for the 60 g kg)1 lipid series and 533 g kg)1 CL and lobster weight that were developed for P. ornatus
DM for the 100 g kg)1 lipid series (Fig. 1). The derived (D. Dennis & T. Skewes, pers. comm.), these CL correspond
dietary protein to energy ratio supporting these maximal to lobster weights of approximately 0.6, 4.6 and 34.0 g,
growth rates was 29.2 and 29.1 g CP MJ)1 GE, respectively. respectively and thus SGRs of 1.7 and 1.5% day)1 for the
These values are very similar to the advocated optimal value weight-increment periods of 0.58–4.6 and 4.6–34 g, respec-
of 30 g CP MJ)1 GE for shrimp (Cuzon & Guillaume 1997). tively. These SGRs are higher than those achieved by lobsters

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236 D.M. Smith et al.

fed the best of the laboratory-pelleted diets in experiment 1 stimulants have on food intake may result in the develop-
(Table 4), but are a little below those for lobsters fed the ment of an experimental diet that produces higher growth
kuruma shrimp diet in experiment 2 (Table 6). However, it rates than achieved in this study, which could be used as a
should be noted that lobsters in Experiment 1 were inten- vehicle to determine nutrient requirements.
tionally fed at a rate below that required for the lobsters to
reach satiation, in order to more clearly define response
Acknowledgements
relationships to dietary protein and lipid content (Glencross
et al. 1999). The sub-satiation feeding strategy is an This study was financially supported by CSIRO Marine
important experimental tool in nutrition studies as is enables Research and the Australian Fisheries Research & Devel-
valid comparisons to be made among the growth responses opment Corporation (FRDC). It formed part of a nationally
resulting from feeding different diets, without the co-ordinated research effort, the Spiny lobster Enhancement
confounding effects of differences in feed intake. and Aquaculture Subprogram, established by the FRDC.
In the parallel study with P. cygnus using the same series of The authors would like to thank Peter Rothlisberg and
diets, Glencross et al. (2001) reported a similar growth rates Malcolm Brown of CSIRO Marine Research and Denis
to those found in this study. Their lobsters grew faster when Poppi of the University of Queensland for their constructive
fed diets containing 500 or 550 g kg)1 CP. However, the critical reviews of the manuscript.
lobsters fed the reference diet of fresh mussel grew more than
twice as fast as lobsters fed the laboratory-pelleted diets. In
References
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30 C, respectively, and fed to excess of a mixture of frozen Bligh, E.G. & Dyer, W.J. (1959) A rapid method of total lipid
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concentration. The results of experiment 1 also provide growth of juvenile ornate rock lobsters, Panulirus ornatus (Fabri-
indicative information on the protein to energy requirements cius, 1798), in Torres Strait, Australia. Mar. Freshwat. Res., 48,
663–670.
of juvenile P. ornatus lobsters. However, a firm conclusion to Glencross, B.D. & Smith, D.M. (1999) The dietary linoleic and
this effect must await confirmation when lobsters are fed linolenic fatty acids requirements of the shrimp Penaeus monodon.
diets that support growth rates higher than those observed in Aquacult. Nutr., 5, 53–63.
Glencross, B., Smith, D.M. & Williams, K.C. (1998) Effect of dietary
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