Trematode behaviours and the perceptual worlds of parasites

There is a great deal of empirical data and theoretical predictions on the patterns and processes of trematode behaviour, particularly in relation to host-finding activities by the free-living stages and site-finding migrations by the parasitic stages within their hosts. Ecological and evolutionary models of trematode life histories often make explicit assumptions about how these organisms must perceive and respond to signals in their worlds as they move from host to host and as they parasitize each host. Nevertheless, it is unclear how natural selection shapes the parasites' behavioural strategies. In addition, at each stage in their life cycle, trematodes are adorned with elaborate sensory organs and possess sophisticated neuromuscular systems, but it is not clear how they use these complex machinery to perceive their worlds. The purpose of this review is to address this question through insights gathered from a century of research on trematode behaviour. Core theoretical assumptions from modern animal behaviour are used to provide the context for this analysis; a key concept is that all animals have unique perceptual worlds that may be inferred from their behaviours. A critical idea is that all animals possess complex patterns of innate behaviour which can be released by extremely specific signals from the environment. The evidence suggests that trematode parasites live in ecologically predictable aquatic and internal host environments where they perceive only small subsets of the total information available from the environment. A general conclusion is that host finding in miracidia and cercaria, and site-finding by trematodes migrating within their definitive hosts, is accomplished through the release of innate patterns of behaviours which are adaptive within the context of conditions in the worm's environment. Examples from empirical studies are used to support the contention that, despite the apparent complexity of their free-living and parasitic environments, the perceptual worlds of trematodes are impoverished, and complex patterns of behaviour may be released by only a few signals in their environment.