Plant secondary compounds as complementary resources: are they always complementary?

Generalist herbivores typically grow better on mixed- than on single-component diets. This response has been attributed to food complementarities that either enhance the utilization of nutrients or dilute the negative impacts of plant secondary compounds (PSC). For instance, when animals choose between foods that contain diverse PSC, they eat more than animals offered a food that contains just one PSC. In addition to their negative impacts on herbivore fitness, recent evidence suggests that at appropriate doses PSC may provide beneficial effects to herbivores (i.e., by reducing parasitic infections). Thus, complementarities among diverse PSC may not only influence an herbivore's ability to consume food but also reduce the incidence of disease. We assessed the complementary effects of two PSC by offering sheep (Ovis aries) a choice of foods containing condensed tannins and saponins while challenged with a parasitic (Haemonchus contortus) infection. Animals offered a choice ate more than animals just offered tannins or saponins in single rations. However, sheep offered choices displayed greater fecal egg counts (an indirect measurement of parasitic burdens) than sheep offered single rations. Thus, saponin- and tannin-containing foods were complementary resources regarding nutrient intake but antagonistic regarding effects on parasitic loads. The nature of the relationship among PSC may depend on the dimension (i.e., nutrient intake, disease) where the interaction occurs. A unifying currency such as growth or reproductive output may help understand the trade-offs between costs (disease) and benefits (nutrient and medicine intake) for herbivores grazing multiple PSC.