“…Depending on biotic and abiotic conditions, natural biofilms can be complex, varying with factors such as the availability of nutrients (Wang et al, ), intensity and wavelength of light (Paterson et al, ), hydrodynamic conditions (Celmer et al, ; Chao et al, ; Kim et al, ; Stoodley et al, ), and grazing pressure (Sommer, ; Wey et al, ). Moreover, while natural sediments provide an excellent substratum for biofilm growth (large surface to volume ratio, rich in nutrients, and porous structure) (Fagherazzi et al, ), colonization is very different from impermeable surfaces (such as biofilms growing on glass slides) since EPS may be distributed throughout the porous medium (Davis et al, ; Gerbersdorf et al, ; Jaiswal et al, ; Van Colen et al, ; Volk et al, ).…”