Modeling plankton mixotrophy: A mechanistic model consistent with the shuter-type biochemical approach

Mixotrophy, i.e., the ability to combine phototrophy and phagotrophy in one organism, is now recognized to be widespread among photic-zone protists and to potentially modify the structure and functioning of planktonic ecosystems. However, few biogeochemical/ecological models explicitly include this mode of nutrition, owing to the large diversity of observed mixotrophic types, the few data allowing the parameterization of physiological processes, and the need to make the addition of mixotrophy into existing ecosystem models as simple as possible. We here propose and discuss a flexible model that depicts the main observed behaviors of mixotrophy in microplankton. A first model version describes constitutive mixotrophy (the organism photosynthesizes by use of its own chloroplasts). This model version offers two possible configurations, allowing the description of constitutive mixotrophs (CMs) that favor either phototrophy or heterotrophy. A second version describes non-constitutive mixotrophy (the organism performs phototrophy by use of chloroplasts acquired from its prey). The model variants were described so as to be consistent with a plankton conceptualization in which the biomass is divided into separate components on the basis of their biochemical function (Shuter-approach; Shuter, 1979). The two model variants of mixotrophy can easily be implemented in ecological models that adopt the Shuter-approach, such as the MIRO model (Lancelot et al., 2005), and address the challenges associated with modeling mixotrophy