Nematode Infections In Ruminants Research Articles

Stochastic Modelling of Environmental Variation for Biological Populations

We examine stochastic effects, in particular environmental variability, in population models of biological systems. Some simple models of environmental stochasticity are suggested, and we demonstrate a number of analytic approximations and simulation-based approaches that can usefully be applied to them. Initially, these techniques, including moment-closure approximations and local linearization, are explored in the context of a simple and relatively tractable process. Our presentation seeks to introduce these techniques to a broad-based audience of applied modellers. Therefore, as a test case, we study a natural stochastic formulation of a non-linear deterministic model for nematode infections in ruminants, proposed by Roberts and Grenfell (1991). This system is particularly suitable for our purposes, since it captures the essence of more complicated formulations of parasite demography and herd immunity found in the literature. We explore two modes of behaviour. In the endemic regime the stochastic dynamic fluctuates widely around the non-zero fixed points of the deterministic model. Enhancement of these fluctuations in the presence of environmental stochasticity can lead to extinction events. Using a simple model of environmental fluctuations we show that the magnitude of this system response reflects not only the variance of environmental noise, but also its autocorrelation structure. In the managed regime host-replacement is modelled via periodic perturbation of the population variables. In the absence of environmental variation stochastic effects are negligible, and we examine the system response to a realistic environmental perturbation based on the effect of micro-climatic fluctuations on the contact rate. The resultant stochastic effects and the relevance of analytic approximations based on simple models of environmental stochasticity are discussed.

Veterinary Parasitology: Developments in Immunology, Epidemiology and Control

1. Preface R. J. Martin 2. Simulating Dictoyocaulus viviparus infection in calves: the parasitic phase H. W. Ploeger and M. Eysker 3. Vaccine development and diagnostics of Dictoyocaulus viviparus J. B. McKeand 4. The immune response and the evaluation of acquired immunity against gastrointestinal nematodes in cattle: a review E. Claerebout and J. Vercruysse 5. Development of vaccines against gastrointestinal nematodes D. Knox 6. Immunological responses of sheep to Haemonchus contortus H. D. F. H. Schallig 7. Impact of nutrition on the pathophysiology of bovine trypanosomiasis P. H. Holmes, E. Katunguka-Rwakishaya, J. J. Benison, G. J. Wassink and J. J. Parkins 8. Electrophysiological investigation of anthelmintic resistance R. J. Martin 9. The development of anthelmintic resistance in sheep nematodes F. Jackson and R. L. Coop 10. Value of present diagnostic methods for gastrointestinal nematode infections in ruminants M. Eysker and H. W. Ploeger 11. Prospects for controlling animal parasitic nematodes by predacious micro fungi M. Larsen 12. Onchocerca ochengi infections in cattle as a model for human onchocerciasis: recent developments A. J. Trees, S. P. Graham, A. Renz, A. E. Bianco and V. Tanya.

Stochastic effects in a model of nematode infection in ruminants

Stochastic effects in a model of nematode infection in ruminants

Stochastic effects in a model of nematode infection in ruminants

We illustrate the importance of stochastic effects in population models of biological systems and demonstrate a number of analytic and simulation-based approaches that can usefully be applied to such models. In so doing, we compare the stochastic approach to the more usual deterministic one. The model studied represents the gastrointestinal infection of ruminants by nematodes when the hosts maintain a fixed density. The incorporation of a feedback mechanism, which accounts for the immune response of the infected animals, results in a highly nonlinear model; similar forms of nonlinearity are a feature of many plausible models in population biology. In the absence of an analytic solution to the full stochastic model we explore a number of approximations and compare them to simulations of the full stochastic process. We explore three modes of behaviour of the system. In the endemic regime the stochastic system fluctuates widely around the non-zero fixed points of the deterministic model. In the managed regime, where the system is subject to external periodic perturbation, stochastic effects are negligible. Finally, we find that in a regime in which the deterministic model predicts the long-term persistence of oscillations the stochastic model shows that extinction can occur. Of the approximation procedures we consider, the Normal approximation to the full stochastic process is the most generally applicable, and it is also the most accurate in the light of simulation results. Local linearization provides reasonably accurate prediction of the variance-covariance structure, and a transfer function approach allows calculation of the time-lagged auto- and cross-correlations in the endemic regime. Linearization of the stochastic updates themselves results in poor prediction of the population variances.

Nutrition and Parasite Interaction

This overview focuses on the interaction between nutritional status and gastrointestinal nematode infection in ruminants and considers: (i) the influence of the parasite on host metabolism; and (ii) the effect of host nutrition on the establishment and survival of parasite populations, the development of the host-immune response and the pathophysiology of infection. Gastrointestinal nematodes reduce voluntary feed intake and efficiency of feed utilisation, a key feature being an increased endogenous loss of protein into the gastrointestinal tract. Overall there is movement of protein from productive processes into repair of the gastrointestinal tract, synthesis of plasma proteins and mucoprotein production. Although reduction in feed intake is a major factor contributing to the reduced performance of parasitised ruminants, the underlying mechanisms of the anorexia are poorly understood. Supplementation of the diet with additional protein does not appear to affect initial establishment of nematode infections but the pathophysiological consequences are generally more severe on lower planes of protein nutrition. The main effect of protein supplementation is to increase the rate of acquisition of immunity and increase resistance to reinfection and this has been associated with an enhanced cellular immune response in the gastrointestinal mucosa. The unresponsiveness of the young lamb can be improved by dietary protein supplementation. Recent trials have shown that growing sheep offered a free choice between a low and a high protein ration are able to modify their diet selection in order to alleviate the increase in protein requirements which result from gastrointestinal nematode infection. Studies on the influence of nutrition on the expression of genotype have shown that the benefits of a superior genotype are not lost on a low protein diet whereas a high protein diet can partially emeliorate the disadvantages of an inferior genotype. In addition to dietary protein both macro-minerals and trace elements can influence the host-parasite relationship.

Nutrition and parasite interaction

This overview focuses on the interaction between nutritional status and gastrointestinal nematode infection in ruminants and considers: (i) the influence of the parasite on host metabolism; and (ii) the effect of host nutrition on the establishment and survival of parasite populations, the development of the host-immune response and the pathophysiology of infection. Gastrointestinal nematodes reduce voluntary feed intake and efficiency of feed utilisation, a key feature being an increased endogenous loss of protein into the gastrointestinal tract. Overall there is movement of protein from productive processes into repair of the gastrointestinal tract, synthesis of plasma proteins and mucoprotein production. Although reduction in feed intake is a major factor contributing to the reduced performance of parasitised ruminants, the underlying mechanisms of the anorexia are poorly understood. Supplementation of the diet with additional protein does not appear to affect initial establishment of nematode infections but the pathophysiological consequences are generally more severe on lower planes of protein nutrition. The main effect of protein supplementation is to increase the rate of acquisition of immunity and increase resistance to reinfection and this has been associated with an enhanced cellular immune response in the gastrointestinal mucosa. The unresponsiveness of the young lamb can be improved by dietary protein supplementation. Recent trials have shown that growing sheep offered a free choice between a low and a high protein ration are able to modify their diet selection in order to alleviate the increase in protein requirements which result from gastrointestinal nematode infection. Studies on the influence of nutrition on the expression of genotype have shown that the benefits of a superior genotype are not lost on a low protein diet whereas a high protein diet can partially emeliorate the disadvantages of an inferior genotype. In addition to dietary protein both macro-minerals and trace elements can influence the host-parasite relationship.

Genetic resistance to parasitic disease: particularly of resistance in ruminants to gastrointestinal nematodes

There is substantial variation among individuals in susceptibility to a wide variety of parasitic diseases and part of this variation in susceptibility is due to genetic factors. The challenge now is to determine the best methods of using the variation to improve our understanding of parasitic infection and to reduce the ravages of parasitic disease. Scientific and commercial applications will depend upon the type of genetic variation. Variation among breeds can be easily exploited by a policy of breed substitution. Variation within a breed can be exploited by selective breeding to improve resistance to infection or to disease, but more work is needed to develop selection indices which are acceptable to livestock breeders. Identifying genes which contribute to the variation in resistance provides a better understanding of the mechanisms of resistance but more work is needed to determine if such genes, alone or in combination, account for a sufficient proportion of the variation in resistance to allow marker assisted selection. A comparison of responses in susceptible and resistant stock provides a powerful tool to distinguish among protective, irrelevant and pathological responses. These themes have been illustrated by three studies of gastrointestinal nematode infections in ruminants.