The effects of herbivorous zooplankton on the sedimentation of particles out of the euphotic zone are examined with mathematical models, a large-scale field experiment, and descriptive data from a eutrophic lake. The theory is rooted in the population dynamics of phytoplankton and zooplankton and so explicitly accounts for the potential effect of zooplankton grazing on primary production and the connection between the rate at which phytoplankton cells sink and sustainable zooplankton biomass. The models predict positive, negative, or unimodal zooplankton effects, depending on the values of four parameters: rate of direct phytoplankton sinking, fraction of zooplankton fecal material exiting the euphotic zone, zooplankton assimilation efficiency, and system productivity. Models were parameterized with data from a eutrophic lake to make a priori predictions about the shape and direction of zooplankton effects. Predictions were tested against the results of an independent experiment in which a gradient of Daphnia biomass was established in large enclosures. Daphnia negatively affected sedimentation rates of carbon, nitrogen, and phosphorus in the enclosure experiment, which confirmed model predictions. Daphnia had a strong negative effect on phytoplankton biomass, and phytoplankton biomass was positively correlated with sedimentation in the enclosures. Experimental results were congruent with relationships between Daphnia biomass and sedimentation rate in the lake. Successful application of the theory suggests that these models may be of utility for assessing the direction of zooplankton effects on vertical flux in other systems. More generally, the models help to identify parameters that should be measured in studies of zooplankton effects on downward particle flux.