Trickle or clumped infection process? An analysis of aggregation in the weights of the parasitic roundworm of humans, Ascaris lumbricoides

Studying the distribution of parasitic helminth body size across a population of definitive hosts can advance our understanding of parasite population biology. Body size is typically correlated with egg production. Consequently, inequalities in body size have been frequently measured to infer variation in reproductive success (VRS). Body size is also related to parasite age (time since entering the definitive host) and potentially provides valuable information on the mode of acquisition and establishment of immature (larval) parasites within the host: whether parasites tend to establish singly or in aggregates. The mode of acquisition of soil-transmitted helminths has been a theoretical consideration in the parasitological literature but has eluded data-driven investigation. In this paper, we analyse individual Ascaris lumbricoides weight data collected from a cohort of human hosts before and after re-infection following curative treatment, and explore its distribution within and among individuals in the population. Lorenz curves and Gini coefficients indicate that levels of weight inequality (a proxy for VRS) in A.lumbricoides are lower than other published estimates from animal-helminth systems. We explore levels of intra-host weight aggregation using statistical models to estimate the intraclass correlation coefficient (ICC) while adjusting for covariates using a flexible fractional polynomial transformation approach capable of handling non-linear functional relationships. The estimated ICCs indicate that weights are aggregated within hosts both at equilibrium and after re-infection, suggesting that parasites may establish within the host in clumps. The implications of a clumped infection process are discussed in terms of ascariasis transmission dynamics, control and anthelmintic resistance.