Modelling growth and reproduction of the Pacific oyster Crassostrea gigas: Advances in the oyster-DEB model through application to a coastal pond
Introduction
Energetic budget models have been widely applied to bivalves in aquaculture, especially for assessing the carrying capacity of coastal systems (e.g. Héral, 1993, Dowd, 1997, Bacher et al., 1998, Duarte et al., 2003, Grant et al., 2003). Such models are based on ecophysiological modelling that details the physiological processes and energetics of an organism in response to environmental fluctuations. Most energetic models of bivalves are net production models (e.g. Ross and Nisbet, 1990, Raillard et al., 1993, Smaal and Widdows, 1994, Barillé et al., 1997, Campbell and Newell, 1998, Grant and Bacher, 1998, Scholten and Smaal, 1998, Ren and Ross, 2001, Hawkins et al., 2002, Gangnery et al., 2003) based on the Scope for Growth (SFG) concept (Bayne and Newell, 1983). Dynamic energy budget (DEB) models are a different type of energetic model that describes the rates at which organisms assimilate and utilise energy for maintenance, growth and reproduction. DEB modelling has also been applied to various bivalves (e.g. Van Haren and Kooijman, 1993, Ren and Ross, 2005, Cardoso et al., 2006, Pouvreau et al., 2006). The DEB theory is based on physical and chemical assumptions for individual energetics (Kooijman, 1986, Kooijman, 2000), whereas the energetics in SFG models are empirically-based using allometric relationships (Lika and Nisbet, 2000, Nisbet et al., 2000, Van der Meer, 2006).
DEB theory has recently been more specifically applied to the Pacific oyster Crassostrea gigas (e.g. Van der Veer and Alunno-Bruscia, 2006). Pouvreau et al. (2006) validated the DEB model for this species reared in various different environments and concluded that the model could be applied in many ecosystems where C. gigas is cultured. Our study aims to refine the initial version of the oyster-DEB model by Pouvreau et al. (2006) and to test the updated version under new environmental conditions in an Atlantic oyster pond. More precisely, this paper describes how effects of temperature on physiological processes have been modified and improved in the model, compared with the extended Arrhenius relationship proposed by Van der Veer et al. (2006). Model simulations were performed using a number of food quantifiers to identify those most suitable for predicting the growth of C. gigas. In the initial oyster-DEB model by Pouvreau et al. (2006), chlorophyll a concentration (chl-a) was the only food quantifier tested. Although chl-a is often used to estimate the phytoplankton biomass available for filter-feeders, there are many sources of discrepancies when using chl-a: (1) quantity of chl-a per phytoplankton cell varies a great deal over the year (Llewellyn et al., 2005), (2) the chl-a measured includes inputs from many sources, e.g. from macroalgae and river detritic particles containing both labile and refractory components, toxic algae, and picoplankton such as Chlorophyta flagellates, which are not retained or assimilated by C. gigas (Barillé et al., 1993, Dupuy et al., 2000b). Moreover, chl-a is a photosynthetic pigment and not a nutritive compound for filter-feeders. We therefore tested additional food quantifiers: particulate organic matter (POM), particulate organic carbon (POC) and, for the first time to our knowledge, phytoplankton enumeration expressed both in cell number per litre and in cumulative biovolume of cells. This assessment aimed to determine the most relevant quantifier to explain oyster growth and reproduction throughout the year.
Section snippets
Model description
A detailed description of the DEB model validated for C. gigas is given in Pouvreau et al. (2006). The general framework of the oyster-DEB model, i.e. the equations and most of the DEB parameters, was kept similar in the present study and hence only a brief summary of the main outline is presented here. The present study, however, focuses on new improvements concerning the temperature effect on oyster physiology and the choice of the most relevant food quantifier.
Forcing variables
Temporal variations in the forcing variables between January 2006 and January 2007 are illustrated in Fig. 3, Fig. 4, for the seawater temperature and food quantifiers, respectively. Seawater temperature showed a classical seasonal pattern from 3 °C to 30 °C between January and July 2006 (Fig. 3). POC varied from 0.2 µg L− 1 in February to 2.7 µg L− 1 in May (Fig. 4A). POM showed a similar pattern to POC, with concentration varying from 2 mg L− 1 in February 2006 to 12 mg L− 1 in January 2007 (Fig.
Discussion
In this study, several parameters of the existing oyster-DEB model developed by Pouvreau et al. (2006) were reconsidered and modified. The resulting second version of the model was then applied and validated on a new dataset of environmental and growth variables. The model was run with different food quantifiers, and phytoplankton enumeration demonstrated its reliability to represent the best the available food explaining observed oyster growth. We first discuss the food quantifier assessment,
Acknowledgements
Y. Bourlès was supported by funding of Région Poitou-Charentes and Ifremer during his PhD project. We would like to thank two anonymous referees for their helpful comments on the manuscript. The members of the European Research Group AquaDEB (http://www.ifremer.fr/aquadeb/) are gratefully acknowledged for the stimulating discussions and useful comments. This paper benefited from helpful comments and English revision by H. McCombie.
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