title: From plants to birds: higher avian predation rates in trees responding to insect herbivory.
abstract: BACKGROUND
  An understanding of the evolution of potential signals from plants to the predators of their herbivores may provide exciting examples of co-evolution among multiple trophic levels. Understanding the mechanism behind the attraction of predators to plants is crucial to conclusions about co-evolution. For example, insectivorous birds are attracted to herbivore-damaged trees without seeing the herbivores or the defoliated parts, but it is not known whether birds use cues from herbivore-damaged plants with a specific adaptation of plants for this purpose.
  
  
  METHODS
  We examined whether signals from damaged trees attract avian predators in the wild and whether birds could use volatile organic compound (VOC) emissions or net photosynthesis of leaves as cues to detect herbivore-rich trees. We conducted a field experiment with mountain birches (Betula pubescens ssp. czerepanovii), their main herbivore (Epirrita autumnata) and insectivorous birds. Half of the trees had herbivore larvae defoliating trees hidden inside branch bags and half had empty bags as controls. We measured predation rate of birds towards artificial larvae on tree branches, and VOC emissions and net photosynthesis of leaves.
  The predation rate was higher in the herbivore trees than in the control trees. This confirms that birds use cues from trees to locate insect-rich trees in the wild. The herbivore trees had decreased photosynthesis and elevated emissions of many VOCs, which suggests that birds could use either one, or both, as cues. There was, however, large variation in how the VOC emission correlated with predation rate. Emissions of (E)-DMNT [(E)-4,8-dimethyl-1,3,7-nonatriene], beta-ocimene and linalool were positively correlated with predation rate, while those of highly inducible green leaf volatiles were not. These three VOCs are also involved in the attraction of insect parasitoids and predatory mites to herbivore-damaged plants, which suggests that plants may not have specific adaptations to signal only to birds.
authors.affiliation: Section of Ecology, Department of Biology, University of Turku, Turku, Finland.
authors.forename: Elina
authors.initials: E
authors.lastname: Mäntylä
authors.forename: Giorgio A
authors.initials: GA
authors.lastname: Alessio
authors.forename: James D
authors.initials: JD
authors.lastname: Blande
authors.forename: Juha
authors.initials: J
authors.lastname: Heijari
authors.forename: Jarmo K
authors.initials: JK
authors.lastname: Holopainen
authors.forename: Toni
authors.initials: T
authors.lastname: Laaksonen
authors.forename: Panu
authors.initials: P
authors.lastname: Piirtola
authors.forename: Tero
authors.initials: T
authors.lastname: Klemola
mesh_terms: D000818:Animals; D001717:Birds / Q000502:physiology*; D004777:Environment; D019143:Evolution, Molecular; D005247:Feeding Behavior; D005387:Finland; D020387:Food Chain; D007313:Insecta; D008954:Models, Biological; D009930:Organic Chemicals / Q000737:chemistry; D010788:Photosynthesis; D018515:Plant Leaves / Q000378:metabolism; D011157:Population Dynamics; D011235:Predatory Behavior; D014197:Trees
chemical_list: D009930:Organic Chemicals