title: High herbivore pressure favors constitutive over induced defense.
abstract: Theoretical and empirical studies show that, when past or current herbivory is a reliable cue of future attack and defenses are costly, defenses can be induced only when needed and thereby permit investment in other functions such as growth or reproduction. Theory also states that, in environments where herbivory is constantly high, constitutive defenses should be favored. Here, we present data to support the second aspect of the induced resistance hypothesis. We examined herbivore-induced responses for four species of Inga (Fabaceae), a common canopy tree in Neotropical forests. We quantified chemical defenses of expanding leaves, including phenolic, saponin and toxic amino acids, in experimental field treatments with and without caterpillars. Because young leaves lack fiber and are higher in protein than mature leaves, they typically lose >25% of their leaf area during the few weeks of expansion. We predicted that the high rates of attack would select for investment in constitutive defenses over induction. Our data show that chemical defenses were quite unresponsive to herbivory. We demonstrated that expanding leaves showed no or only small increases in investment in secondary metabolites, and no qualitative changes in the phenolic compound profile in response to herbivory. The proteinogenic amino acid tyrosine, which can be toxic at high concentrations, showed the greatest levels of induction.
  
  
  
  These results provide some of the first support for theoretical predictions that the evolution of induced vs. constitutive defenses depends on the risk of herbivory. In habitats with constant and high potential losses to herbivores, such as tropical rainforests, high investments in constitutive defenses are favored over induction.
authors.affiliation: Department of Biology University of Utah 257S 1400E Salt Lake City Utah 84112.
authors.forename: Ryan J
authors.initials: RJ
authors.lastname: Bixenmann
authors.affiliation: Department of Biology University of Utah 257S 1400E Salt Lake City Utah 84112; Smithsonian Tropical Research Institute Box 0843-03092 Balboa Republic of Panama.
authors.forename: Phyllis D
authors.initials: PD
authors.lastname: Coley
authors.affiliation: Department of BiologyUniversity of Utah 257S 1400E Salt Lake City Utah 84112; German Centre for Integrative Biodiversity Research (iDiv), Halle-Jena-Leipzig Deutscher Platz 5e Leipzig 04103 Germany.
authors.forename: Alexander
authors.initials: A
authors.lastname: Weinhold
authors.affiliation: Department of Biology University of Utah 257S 1400E Salt Lake City Utah 84112; Smithsonian Tropical Research Institute Box 0843-03092 Balboa Republic of Panama.
authors.forename: Thomas A
authors.initials: TA
authors.lastname: Kursar
keywords: Defense chemistry; induced defenses; phenolics; plant–herbivore interactions; saponins; secondary metabolites; tropical forest