Random, top-down, or bottom-up coexistence of parasites: malaria population dynamics in multi-parasitic settings.

Epidemiological models concerned with the control of malaria using interventions such as bed nets and vaccines increasingly incorporate realistic aspects of malaria biology. The increasing complexity of these models limits their ability to abstract ecological processes and to address questions on the regulation of population dynamics using time-series data, particularly in regards to interactions between different pathogens and the regulatory role of innate (bottom-up) and acquired (top-down) immunity. We use a theoretical framework to test hypotheses on the importance of population-level immunity and parasite abundance in regulating the population dynamics of malaria. We use qualitative loop analyses to examine the sign of the interaction between Plasmodium falciparum and P. vivax at the population level, and we discuss implications of this sign for the within-host regulation of parasites. Our analyses of monthly malaria time-series data from the island of Espirito Santo, Vanuatu (1983-1997), show that the dynamics of P. falciparum are not sensitive to P. vivax, whereas infections by the latter increase in response to those of the former. These results support a differential use of resources inside the hosts, a resource-consumer interaction between hosts and their immune system, and within-host regulation of parasites. Finally, our results emphasize the need to better understand factors regulating malaria dynamics before developing control strategies and call for the use of control strategies directed at the interruption of transmission, such as vector control and the use of bed nets.