A model for nematode locomotion in soil
Locomotion of nematodes in soil is important for both practical and theoretical reasons. We constructed a model for rate of locomotion. The first model component is a simple simulation of nematode movement among finite cells by both random and directed behaviours. Optimisation procedures were used to fit the simulation output to data from published experiments on movement along columns of soil or washed sand, and thus to estimate the values of the model's movement coefficients. The coefficients then provided an objective means to compare rates of locomotion among studies done under different experimental conditions. The second component of the model is an equation to predict the movement coefficients as a function of controlling factors that have been addressed experimentally: soil texture, bulk density, water potential, temperature, trophic group of nematode, presence of an attractant or physical gradient and the duration of the experiment. Parameters of the equation were estimated by optimisation to achieve a good fit to the estimated movement coefficients. Bulk density, which has been reported in a minority of published studies, is predicted to have an important effect on rate of locomotion, at least in fine-textured soils. Soil sieving, which appears to be a universal practice in laboratory studies of nematode movement, is predicted to negatively affect locomotion. Slower movement in finer textured soils would be expected to increase isolation among local populations, and thus to promote species richness. Future additions to the model that might improve its utility include representing heterogeneity within populations in rate of movement, development of gradients of chemical attractants, trade-offs between random and directed components of movement, species differences in optimal temperature and water potential, and interactions among factors controlling locomotion.