Growth and energetics in the stickleback-Schistocephalus host-parasite system: a review of experimental infection studies

Three-spined sticklebacks in natural lacustrine populations are often infected with plerocercoids of the indirectly transmitted pseudophyllidean cestode Schistocephalus solidus. Field studies typically show infections to be associated with reduced host condition, gonadogenesis and energy reserves, though infection phenotypes can vary considerably both between and within host populations. Experimental infection studies allow the impact of infections on hosts to be studied under a variety of rearing conditions, and so can be used to determine the environmental component of infection phenotypes. Here, we review recent laboratory studies undertaken by our group, examining the growth and condition of experimentally infected fish reared under conditions that differed in terms of absolute ration, temporal pattern of feeding and level of competition between fish. We compare infection phenotypes generated in our experimental studies with those of fish sampled in field based studies. Experimental studies in which infected fish were reared under competition for limited food resources, or were fed a restricted diet, generated infection phenotypes that most closely resembled those found in the majority of natural populations. When access to food was unrestricted, however, infected fish were able to sustain high growth rates and lay down energy reserves. If experimental studies are to be used to understand the impact of infection under natural conditions, husbandry protocols that closely match field conditions must be designed. We suggest that a full understanding of the impact of parasites on their hosts can only be gained by integrating controlled laboratory experiments with detailed field studies. The stickleback-Schistocephalus system is ideally suited to examining these questions, and we provide several suggestions for future research.