Review: Herbivory and the power of phytochemical diversity on animal health.

Plant secondary compounds (PSCs) were thought to be waste products of plant metabolism when first identified in the mid-1800 s. Since then, many different roles have been recognized for these chemicals. With regard to their function as defense, PSCs can negatively impact different cellular and metabolic processes in the herbivore, causing illness and reductions in feed intake. This penalty on fitness also applies to other trophic levels, like the microorganisms and parasites that infect herbivores and thus, PSCs at certain doses may function as medicines. In turn, herbivores evolved learning mechanisms to cope with the constant variability in their environment and physiological needs. Under this context, foraging can be viewed as the quest for substances in the external environment that provide homeostatic utility to the animal. For instance, herbivores increase preference for PSC-containing feeds that negatively impact infectious agents (i.e., therapeutic self-medication). Given that some classes of PSCs like polyphenols present antioxidant, antiinflammatory, immunomodulatory and prebiotic properties, chronic and sustained consumption of these chemicals results in robust animals that are tolerant to disease (i.e., prophylactic self-medication). Foraging plasticity in terms of the quality and quantity of nutrients ingested in the absence and during sickness may also influence immunocompetence, resistance and resilience to infection, and thus can be interpreted as another form of medication. Finally, self-medicative behaviors can be transmitted through social learning. We suggest that foraging studies will benefit from exploring self-medicative behaviors in chemically diverse plant communities, in particular when considering the vast diversity of PSC structures (more than 200 000) observed in nature. We then lay out a framework for enhancing the medicinal effects of PSCs on grazing herbivores. We propose landscape interventions through the establishment of resource patches or "islands" with a diversity of PSC-containing forages (e.g., legumes, herbs, shrubs) in monotonous rangelands or pasturelands, viewed as a "sea" of low-diversity vegetation devoid of functional biochemicals. Strategies aimed at enhancing the diversity of plant communities lead to heterogeneity in chemical, structural and functional landscape traits that offer options to foragers, and thus allow for balanced diets that maintain and restore health. Beyond animal health, such heterogeneity promotes a broad array of ecosystem services that significantly improve landscape resilience to environmental disturbances.