SPEAD 1.0 – Simulating Plankton Evolution with Adaptive Dynamics in a two-trait continuous fitness landscape applied to the Sargasso Sea

摘要

Diversity plays a key role in the adaptive capacity of marine ecosystems to environmental changes. However, modelling the adaptive dynamics of phytoplankton traits remains challenging due to the competitive exclusion of sub-optimal phenotypes and the complexity of evolutionary processes leading to optimal phenotypes. Trait diffusion (TD) is a recently developed approach to sustain diversity in plankton models by introducing mutations, therefore allowing the adaptive evolution of functional traits to occur at ecological timescales. In this study, we present a model called Simulating Plankton Evolution with Adaptive Dynamics (SPEAD) that resolves the eco-evolutionary processes of a multi-trait plankton community. The SPEAD model can be used to evaluate plankton adaptation to environmental changes at different timescales or address ecological issues affected by adaptive evolution. Phytoplankton phenotypes in SPEAD are characterized by two traits, the nitrogen half-saturation constant and optimal temperature, which can mutate at each generation using the TD mechanism. SPEAD does not resolve the different phenotypes as discrete entities, instead computing six aggregate properties: total phytoplankton biomass, the mean value of each trait, trait variances, and the inter-trait covariance of a single population in a continuous trait space. Therefore, SPEAD resolves the dynamics of the population's continuous trait distribution by solving its statistical moments, wherein the variances of trait values represent the diversity of ecotypes. The ecological model is coupled to a vertically resolved (1D) physical environment, and therefore the adaptive dynamics of the simulated phytoplankton population are driven by seasonal variations in vertical mixing, nutrient concentration, water temperature, and solar irradiance. The simulated bulk properties are validated by observations from Bermuda Atlantic Time-series Studies (BATS) in the Sargasso Sea. We find that moderate mutation rates sustain trait diversity at decadal timescales and soften the almost total inter-trait correlation induced by the environment alone, without reducing the annual primary production or promoting permanently maladapted phenotypes, as occur with high mutation rates. As a way to evaluate the performance of the continuous trait approximation, we also compare the solutions of SPEAD to the solutions of a classical discrete entities approach, with both approaches including TD as a mechanism to sustain trait variance. We only find minor discrepancies between the continuous model SPEAD and the discrete model, with the computational cost of SPEAD being lower by 2?orders of magnitude. Therefore, SPEAD should be an ideal eco-evolutionary plankton model to be coupled to a general circulation model (GCM) of the global ocean.