After the exponential growth phase, variability in the scattering efficiency of phytoplankton cells over their complete life cycle is not well characterised. Bulk measurements are impacted by senescent cells and detritrus. Thus the analysis of the evolution of the optical properties thanks to their morphological and/or intra-cellular variations remains poorly studied. Using the Cytosense flow cytometer (CytoBuoy b.v., NL), the temporal course of the forward and sideward efficiencies of two phytoplankton species (Thalassiosira pseudonana and Chlamydomonas concordia) were analyzed during a complete life-cycle. These two species differ considerably from a morphological point of view. Over the whole experiment, the forward and sideward efficiencies of Thalassiosira pseudonana were, on average, respectively 2.2 and 1.6 times higher than the efficiencies of Chlamydomonas concordia. Large intra-species variability of the efficiencies were observed over the life cycle of the considered species. It highlights the importance of considering the optical properties of phytoplankton cells as a function of the population growth stage of the considered species. Furthermore, flow cytometry measurements were combined with radiative transfer simulations and biogeochemical and optical measurements. Results showed that the real refractive index of the chloroplast is a key parameter driving the sideward signal and that a simplistic two-layered model (cytoplasm-chloroplast) seems particularly appropriate to represent the phytoplankton cells.

The bulk backscattering ratio ( $\widetilde{b_{bp}}$ ) is commonly used as a descriptor of the bulk real refractive index of the particulate assemblage in natural waters. Based on numerical simulations, we analyze the impact of modeled structural heterogeneity of phytoplankton cells on $\widetilde{b_{bp}}$ . $\widetilde{b_{bp}}$ is modeled considering viruses, heterotrophic bacteria, phytoplankton, organic detritus, and minerals. Three case studies are defined according to the relative abundance of the components. Two case studies represent typical situations in open ocean, oligotrophic waters, and phytoplankton bloom. The third case study is typical of coastal waters with the presence of minerals. Phytoplankton cells are modeled by a two-layered spherical geometry representing a chloroplast surrounding the cytoplasm. The $\widetilde{b_{bp}}$ values are higher when structural heterogeneity is considered because the contribution of coated spheres to light backscattering is higher than homogeneous spheres. The impact of heterogeneity is; however, strongly conditioned by the hyperbolic slope $\xi$ of the particle size distribution. Even if the relative abundance of phytoplankton is small (<1%), $\widetilde{b_{bp}}$ increases by about 58% (for $\xi =4$ and for oligotrophic waters), when the heterogeneity is taken into account, in comparison with a particulate population composed only of homogeneous spheres. As expected, heterogeneity has a much smaller impact (about 12% for $\xi =4$ ) on $\widetilde{b_{bp}}$ in the presence of suspended minerals, whose increased light scattering overwhelms that of phytoplankton.

The bulk backscattering ratio ($\tilde{b_{bp}}$) is commonly used as a descriptor of the bulk real refractive index of the particulate assemblage in natural waters. Based on numerical simulations, we analyze the impact of heterogeneity of phytoplankton cells on $\tilde{b_{bp}}$. $\tilde{b_{bp}}$ is modeled considering viruses, heterotrophic bacteria, phytoplankton, detritus, and minerals. Three study cases are defined according to the relative abundance of these different components. Two study cases represent typical situations in open ocean, outside (No-B/No-M) and inside bloom (B/No-M). The third study case is typical of coastal waters with the presence of minerals. Phytoplankton cells are modeled by a two-layered spherical geometry representing a chloroplast surrounding the cytoplasm. The $\tilde{b_{bp}}$ values are higher when heterogeneity is considered because the contribution of coated spheres to backscattering is higher than homogeneous spheres. The impact of heterogeneity is however strongly conditioned by the hyperbolic slope $\xi$ of the particle size distribution. Even if the relative concentration of phytoplankton is small (<1%), $\tilde{b_{bp}}$ increases by about 60% (for $\xi=4.3$ and for the No-B/No-M water body), when the heterogeneity is taken into account, in comparison with a particulate population only composed by homogeneous spheres. As expected, heterogeneity has a much smaller impact (about 5$\%$ for $\xi=4.3$) on $\tilde{b_{bp}}$ when minerals are added.