title: Density-dependent effects on the weight of female Ascaris lumbricoides infections of humans and its impact on patterns of egg production.
abstract: BACKGROUND
  Ascaris lumbricoides exhibits density-dependent egg production, a process which has a marked impact on both the transmission dynamics and the stability of the parasite population. Evidence suggests that the egg production of female Ascaris is also associated with the size of the worm. If worm size is mediated by density-dependent processes then the size of female worms may have a causal impact upon patterns of Ascaris egg production.
  
  
  RESULTS
  We analyse data collected from a cohort of human hosts, and demonstrate that the per host mean weight (a proxy for size) of female Ascaris is dependent on the number of infecting females (worm burden) following a pattern of initial facilitation followed by limitation. Applying a negative binomial (NB) generalized linear model (GLM) and a zero-inflated negative binomial (ZINB) model we confirm that the per host female mean weight is significantly associated with per host egg production. Despite these associations, the mean weight of female Ascaris has little causal impact on patterns of density-dependent egg output. The ZINB model is able to account for the disproportionately large number of zero egg counts within the data and is shown to be a consistently better fit than the NB model. The probability of observing a zero egg count is demonstrated as being negatively associated with both female worm burden and female mean weight.
  
  
  CONCLUSIONS
  The mean weight of female Ascaris is statistically significantly associated with egg output, and follows a consistent pattern of facilitation preceding limitation with increasing female worm burden. Despite these relationships, incorporation of female Ascaris mean weight into models of egg output has little effect on patterns of density dependence. The ZINB model is a superior fit to the data than the NB model and provides additional information regarding the mechanisms that result in a zero egg count. The ZINB model is shown to be a useful tool for the analysis of individual-based egg output data.
authors.affiliation: Department of Infectious Disease Epidemiology, Faculty of Medicine, St Mary's Campus, Imperial College London, Norfolk Place, London, W2 1PG, UK. m.walker06@imperial.ac.uk.
authors.forename: Martin
authors.initials: M
authors.lastname: Walker
authors.forename: Andrew
authors.initials: A
authors.lastname: Hall
authors.forename: Roy M
authors.initials: RM
authors.lastname: Anderson
authors.forename: María-Gloria
authors.initials: MG
authors.lastname: Basáñez