Particle association of Enterococcus sp. increases growth rates and simulated persistence in water columns of varying light attenuation and turbulent diffusivity.

Predicting water quality and the human health risks associated with sewage-derived microbes requires understanding the fate and transport of these contaminants. Sewage-derived pathogen risks are typically assessed and monitored by measuring concentrations of fecal indicating bacteria (FIB), like Enterococcus sp. Previous research demonstrated that a high fraction of FIB is particle-associated, which can alter FIB dynamics within secondary water bodies. In this study, we experimentally quantified the effect of particle association on dark, temperature- and light-dependent growth and sinking rates of enterococci. Particle association significantly increased dark growth rates, light-dependent growth rates (i.e. decreased mortality), and sinking rates, relative to free-living enterococci. Simulations using a novel, 1-dimensional model parameterized by these rates indicate greater persistence (T) for particle-associated enterococci in water bodies across a wide range of diffuse attenuation coefficients of light (K) and turbulent diffusivity (D) values. In addition, persistence of both fractions increased in simulated turbid and turbulent waters, compared to clear and/or quiescent conditions. Simulated persistence of both fractions also increased when enterococci discharges occurred later in a diel cycle (towards sunset, as opposed to sunrise), especially for the free-living population, because later discharges under our model conditions allowed both fractions to mix deeper before inactivation via sunlight. Model sensitivity testing revealed that T variability was greatest when dark growth rates were altered, suggesting that future empirical studies should focus on quantifying these rates for free-living and particle-associated sewage-derived microbes. Despite greater sensitivity of T to variability in dark growth rates, omitting light-dependent growth rates from simulations dramatically influenced T values. Our results demonstrate that particle association can increase enterococci persistence in receiving waters and highlight the importance of incorporating particle association in future water quality models.