Elsevier

Aquaculture

Volume 499, 15 January 2019, Pages 128-139
Aquaculture

The influence of flesh ingredients format and krill meal on growth and feeding behaviour of juvenile tropical spiny lobster Panulirus ornatus

https://doi.org/10.1016/j.aquaculture.2018.09.019Get rights and content

Highlights

  • This study provided essential information for the inclusion of krill meal and flesh ingredients in feeds for juvenile lobster.

  • Time-series photography analysis was used to better understand the feeding behaviour of early juvenile tropical lobster.

  • This study reports the first finding on the growth performance of P. ornatus juveniles that were hatchery reared from eggs.

Abstract

One of the main challenges for spiny lobster aquaculture is the successful development of formulated feeds that are attractive, readily consumed by lobsters and promote optimal growth and survival. In a 54-day growth trial, we investigated the performance of four moist formulated feeds containing A) non-homogenised flesh ingredients; B) fish meal only; C) 10% krill meal; or D) homogenised flesh ingredients; and a reference diet of shucked blue mussel (Mytilus galloprovincialis) flesh, on growth, nutritional condition and feeding behaviour of juvenile tropical spiny lobster (Panulirus ornatus) reared either communally or individually. Growth and survival achieved by lobsters fed mussel were significantly higher than all other treatments. However, lobsters fed the formulated feed containing 10% krill meal (treatment C) and reared communally had a significantly higher growth, survival and feeding performance when compared to the formulated feeds with inclusion of non-homogenised and homogenised flesh ingredients (treatments A and D). The highest levels of feed interaction and pellet consumption among the formulated feeds were also recorded for treatment C. We demonstrated that the inclusion of krill meal in formulated feeds provides a benefit when compared to the inclusion of the flesh ingredients. Flesh ingredient format (homogenised or non-homogenised) had no effect on growth performance of lobsters, suggesting that the process of homogenisation of flesh ingredients does not provide any benefit in promoting feed consumption. Furthermore, time-series photography analysis trials showed that all the formulated feeds only promoted a feeding response within the first 2–3 h post-feeding, whereas mussel retained its attractiveness throughout the feeding period. These results suggest that the prolonged attractiveness remains an impediment for formulated feed performance in lobster culture. The present study also showed that growth performance and feeding response were higher in lobsters reared communally than individually. As observed in other spiny lobster species, it is likely that social interactions in communal housing may have provided cues that stimulated feeding responses and promoted higher growth rates.

Introduction

Spiny lobsters (Crustacea: Decapoda: Palinuridae) are one of the world's most valuable seafood commodities, forming the basis for major fisheries in numerous countries from tropical to temperate zones (Jeffs, 2010; Phillips and Matsuda, 2011). In 2014, the 306 thousand tonne global production of spiny lobster was valued at more than US $2.7 billion (FAO, 2016). Due to the full exploitation of the fishery in Australia and the capacity of many other world fisheries being exceeded, aquaculture production will become fundamental to meet increasing market demand for spiny lobsters (Dennis et al., 2004; Ehrhardt and Fitchett, 2010; Linnane et al., 2010; Smith, 2017). Over the past decade, studies in larviculture have greatly enhanced the likelihood of sustainable aquaculture of spiny lobsters (Fitzgibbon and Battaglene, 2012; Fitzgibbon et al., 2012, Fitzgibbon et al., 2014, Fitzgibbon et al., 2015; Francis et al., 2014; Jason et al., 2008; Jensen et al., 2013a, Jensen et al., 2013b; Ventura et al., 2015). However, until commercial hatcheries of spiny lobster can be established, commercial aquaculture will continue to be based on collection of wild seed-stock (Jones, 2010; Jones et al., 2015; Petersen and Truong, 2010; Williams, 2004).

Panulirus ornatus is widely considered the best candidate for intensive closed-cycle aquaculture (Phillips and Matsuda, 2011; Phillips, 2006). Panulirus ornatus has high commercial value and demand, a relatively short larval phase (4–6 months) and a rapid growth rate, being able to reach 1 kg after 18 months post-settlement (Jeffs, 2010; Phillips et al., 1992; Phillips, 2006). Although early juveniles P. ornatus are highly cannibalistic, this trait decreases with age and older juveniles can be cultured at high densities with little aggression (Dennis et al., 1997; Jones et al., 2001). Several studies have also shown P. ornatus broad tolerance to environmental conditions including temperature, salinity, turbidity and other water quality parameters (Jones and Shanks, 2009; Jones, 2009; Phillips and Matsuda, 2011).

To date the largest aquaculture production of P. ornatus occurs in Vietnam, based on wild caught seed stock (Petersen and Truong, 2010; Petersen and Phuong, 2011). More recently, other countries in this region, particularly Indonesia, have adopted this practice (Jones et al., 2015). A major concern threatening the existing lobster aquaculture industry has been the reliance on trash fish (low value fish, mollusc and crustaceans) as a source of feed (Jones, 2010; Williams, 2004, Williams, 2007). Several well-identified issues make this feeding practice environmentally unsustainable and inefficient including: strong harvesting pressure on trash fish species; poor feed conversion rate of up to 20:1 (Jones, 2010); inefficient feeding and feed wastage leading to environmental degradation (Nguyen and Dao, 2009); variability in supply and nutritional profile; and difficulties of storage and spoilage (Jones et al., 2015; Nunan et al., 2010). In 2007, the production of spiny lobster in Vietnam declined from 1900 to 1400 metric tonnes due to an outbreak of milky haemolymph syndrome (MHS) associated with poor water quality following bacterial decomposition of trash fish feeds (Hung et al., 2010; Nunan et al., 2010; Petersen and Phuong, 2011; Shields, 2011). The progression to formulated feed would provide several benefits such as nutritional optimisation, ease of usage and storage, flexibility with feeding strategies, improved environmental sustainability and minimisation of waste (Perera and Simon, 2015).

Several studies have focussed on the importance of flesh ingredients and krill meal in P. ornatus feeds with conflicting conclusions. It was hypothesised that the presence of flesh ingredients in lobster feeds is beneficial for promoting improved survival and growth as it increases feed intake (Irvin and Shanks, 2015). Indeed, flesh ingredients not only contain high levels of nutrients but also are texturally softer which may suit the underdeveloped mouthparts of juvenile lobsters (Irvin and Shanks, 2015). In contrast, Irvin and Williams (2009a) reported that sub-adult P. ornatus (> 600 g) had no requirement for flesh ingredients in formulated feeds, which suggests that there may be ontogenetic shifts in formulated feed requirement for P. ornatus. Furthermore, it remains unclear whether flesh feed ingredients encourage feeding through the presence of chemo-attractants (Jones et al., 2015). By investigating the influence of flesh ingredient format (homogenised versus non-homogenised) the present study aims to understand the importance of feed texture on the performance of cultured lobsters.

Krill meal is considered one of the preferred protein sources for feed manufacture of carnivorous crustacean species (Williams, 2007, Williams, 2009). This nutrient-rich additive contains high concentrations of carotenoid pigments, similar amino and fatty acid profile to the cultured lobsters own profile and it enhances the attractiveness and palatability of the feed through the provision of additional free amino acids and peptides (Floreto et al., 2001). In several studies, krill meal and hydrolysates were considered key factors for significant improvements of growth rate in P. ornatus and Panulirus argus (Barclay et al., 2006; Cox and Davis, 2009; Smith et al., 2005). Irvin and Williams (2009a) found that the inclusion of 8% krill hydrolysate in the feeds of sub-adults P. ornatus was necessary to maintain high productivity. It was also hypothesised that krill product can replace the use of flesh ingredients since it sustains lobster's appetite, improving growth. In contrast, krill meal inclusion in formulated feeds of early juveniles resulted in a 44% growth reduction which again suggests ontogenetic differences in P. ornatus feed requirements (Irvin and Shanks, 2015). Considering discrepancies in the performance of formulated feed containing krill meal, further work is required to determine its effectiveness as a growth promoter and attractant, particularly for early juvenile P. ornatus.

Observational studies can provide valuable insights of lobster cryptic behaviour and complex feeding strategies (Carter et al., 2014; Phillips, 2006; Thomas et al., 2003). Spiny lobsters are nocturnal animals, mostly foraging and feeding after dusk to minimise the risk of being predated (Phillips, 2006). Differently from most fish species, which use their vision to locate potential prey, lobsters rely on chemoreception (Phillips, 2006). Several experiments have been conducted to analyse feed components that act as chemo-attractants. Free amino acids (e.g. taurine, glycine), nucleotide adenosine 5-triphosphate and betaine have shown to elicit strong feeding responses in spiny lobsters (Tolomei et al., 2003; Williams, 2007; Williams et al., 2005). However, these feed-attractants rapidly leach from formulated feeds upon immersion in water (Williams et al., 2005) and thus, after 1–2 h, spiny lobsters tend to refuse formulated feeds, whereas, shucked mussel remain attractive for several hours (> 10 h) (Tolomei et al., 2003). Currently, short-time attractiveness remains the major bottleneck for the successful formulation of feeds for spiny lobsters (Phillips and Matsuda, 2011). The combination of the correct physical characteristics (size, shape, stability and buoyancy), as well as attractiveness, is a key factor for the manufacture of formulated feeds (Phillips, 2006). Spiny lobsters are slow, messy feeders and when the physical characteristics of pellets are not adequate, lobster handling and external maceration of the feed before ingestion, can generate considerable food wastage (Sheppard et al., 2002).

The aim of this study was to measure the effects of four moist formulated feeds and a reference diet of shucked blue mussel flesh on growth and nutritional condition of juvenile P. ornatus. These formulated feeds vary in terms of inclusion of flesh ingredients, either homogenised or not, fish meal only or 10% krill meal inclusion. Homogenisation of the flesh ingredients was examined to decipher the influence of texture on feed performance. Krill meal was examined to investigate the apparent contradiction in the literature about the benefits of krill for early juvenile P. ornatus. The present study also reports the first finding on the growth performance of P. ornatus juveniles that were hatchery reared from eggs.

Section snippets

Experimental treatments

Formulated feeds were made at the CSIRO Marine Research Laboratory at Bribie Island, Australia and were based on the Australian Centre for International Agricultural Research (ACIAR) moist feed formulation (Jones et al., 2015). The four formulated feeds, which were formulated to be isonitrogenous (~53% DM) and isoenergetic (~19 MJ kg−1), vary from the ACIAR base feed in terms of inclusion of flesh ingredients, either homogenised or not, fish meal only or 10% krill meal inclusion (Table 1). For

Growth and survival

Lobster initial WW was not significantly different at stocking across all treatments (Table 2). For lobster final WW and WW gain, the interactive effect between rearing condition and feed treatment was significant. Pairwise comparison of the means showed that mussel fed lobsters reared communally had a significant higher final WW than all other treatments and mussel fed lobsters reared individually had a significantly higher final WW than lobsters fed treatment D reared either individually or

Discussion

The results of growth, tissue composition, and time series photography analysis in this study provided important insights into the feed performance and behaviour of juvenile P. ornatus. In both communal and individual culture, lobsters fed with the reference diet of blue mussel achieved higher values of SGR, number of moulting events, survival and biomass than lobsters fed with the formulated feeds (Table 2). Lobsters fed mussel also interacted with the feed longer than lobsters fed the

Conclusion

This study has improved our understanding of the nutritional preferences of early juvenile P. ornatus providing essential information for the development of formulated feeds that are attractive, and optimal for growth and survival. Although the best growth and survival rate were achieved by lobsters fed with blue mussel, encouraging results were obtained by lobsters fed the formulated feed with inclusion of 10% krill meal (treatment C). The results of the time-series photography analysis also

Acknowledgments

Rock Lobster propagation research at IMAS is supported by the Australian Research Council Industrial Transformation Research Hub (project number IH12100032). We thank the staff at the ARC ITH for systems support and assistance in the culture of animals. We thank Mr. Simon Irvin who produced the formulated feeds that were tested in this study. We would also like to acknowledge the input received from the ACIAR project FIS/2014/059 “Expanding spiny lobster aquaculture in Indonesia”. We also thank

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