Parasite-modified behaviour in non-trophic transmission: trematode parasitism increases the attraction between snail intermediate hosts

Many parasites with complex life cycles cause host behavioural changes that increase the likelihood of transmission to the next host. Parasite modification is often found in trophic transmission, but its influence on non-trophic transmission is unclear. In trematodes, transmission from the first to second intermediate host is non-trophic, suggesting that free-swimming larvae (cercariae) emerging in closer proximity to the next host would have higher transmission success. We performed a series of behavioural experiments with echinostome trematodes and their snail hosts to determine if potential second hosts (ramshorn snail, genus Planorbella Haldeman, 1842) were more attracted to parasitized first hosts (marsh pondsnail, Lymnaea elodes Say, 1821). In a Y maze, a responding snail (Planorbella sp.) was placed in the base and its response to five treatments was assessed: no stimulus, turion duckweed (Lemna turionifera Landolt; a food item), non-parasitized L. elodes, parasitized L. elodes, and finally parasitized versus non-parasitized L. elodes. Snails showed some attraction to uninfected snails, but had a stronger response to infected first host snails. These results indicate that potential second host snails were more attracted to parasitized, heterospecific first host snails over non-parasitized heterospecific snails. This study demonstrates that echinostome trematodes alter snail behaviour by changing navigational choices in uninfected potential hosts through a chemical communication mechanism.

Life-history plasticity in hosts (Lymnaea elodes) exposed to differing resources and parasitism

Assessing phenotypic expression across environments is essential for understanding the evolution of life histories, yet relatively few studies have empirically determined the role that multiple environmental factors play in altering animal phenotypes. We used a multifactorial approach to investigate the effects of both infection of Echinostoma revolutum (Frölich, 1802) and nutrient availability on phenotypic expressions in lines of the snail Lymnaea elodes (Say, 1821). Lines were initially established via selfing events followed by breeding (within lines) over the next four generations. Juveniles from each line were then size-matched and randomly exposed to parasite (exposed / sham-exposed) and diet (high protein / low protein) treatments, generating a 2 (line) × 2 (diet) × 2 (exposure) factorial design. Snail growth, reproduction, and survival were monitored over 5 weeks. Analyses revealed an interactive effect of host line and infection status on host growth. Main effects of both snail line and diet also significantly influenced host growth. Reproductive patterns differed between lines, with snails from one line producing egg masses and eggs in all treatments, and snails from the second line producing minimal eggs in only a single treatment. Snail survival remained similar between snail lines. Results from this study suggest that snail life-history traits can vary dramatically as a result of host genetics, the environment, and the interaction between these factors. Reasons for the occurrence and maintenance of this variability in life-history traits are discussed.