title: The timing of herbivore-induced volatile emission in black poplar (Populus nigra) and the influence of herbivore age and identity affect the value of individual volatiles as cues for herbivore enemies.
abstract: BACKGROUND
  The role of herbivore-induced plant volatiles as signals mediating the attraction of herbivore enemies is a well-known phenomenon. Studies with short-lived herbaceous plant species have shown that various biotic and abiotic factors can strongly affect the quantity, composition and timing of volatile emission dynamics. However, there is little knowledge on how these factors influence the volatile emission of long-lived woody perennials. The aim of this study was to investigate the temporal dynamics of herbivore-induced volatile emission of black poplar (Populus nigra) through several day-night cycles following the onset of herbivory. We also determined the influence of different herbivore species, caterpillars of the gypsy moth (Lymantria dispar) and poplar hawkmoth (Laothoe populi), and different herbivore developmental stages on emission.
  
  
  RESULTS
  The emission dynamics of major groups of volatile compounds differed strikingly in response to the timing of herbivory and the day-night cycle. The emission of aldoximes, salicyl aldehyde, and to a lesser extent, green leaf volatiles began shortly after herbivore attack and ceased quickly after herbivore removal, irrespective of the day-night cycle. However, the emission of most terpenes showed a more delayed reaction to the start and end of herbivory, and emission was significantly greater during the day compared to the night. The identity of the caterpillar species caused only slight changes in emission, but variation in developmental stage had a strong impact on volatile emission with early instar L. dispar inducing more nitrogenous volatiles and terpenoids than late instar caterpillars of the same species.
  
  
  CONCLUSIONS
  The results indicate that only a few of the many herbivore-induced black poplar volatiles are released in tight correlation with the timing of herbivory. These may represent the most reliable cues for herbivore enemies and, interestingly, have been shown in a recent study to be the best attractants for an herbivore enemy that parasitizes gypsy moth larvae feeding on black poplar.
authors.forename: Andrea Clavijo
authors.initials: AC
authors.lastname: McCormick
authors.forename: G Andreas
authors.initials: GA
authors.lastname: Boeckler
authors.forename: Tobias G
authors.initials: TG
authors.lastname: Köllner
authors.forename: Jonathan
authors.initials: J
authors.lastname: Gershenzon
authors.forename: Sybille B
authors.initials: SB
authors.lastname: Unsicker
mesh_terms: D000818:Animals; D002940:Circadian Rhythm; D003463:Cues; D060434:Herbivory; D007814:Larva; D009036:Moths / Q000502:physiology*; D018515:Plant Leaves / Q000737:chemistry / Q000502:physiology; D032107:Populus / Q000737:chemistry* / Q000276:immunology / Q000502:physiology; D015398:Signal Transduction; D013729:Terpenes / Q000302:isolation & purification / Q000378:metabolism; D014197:Trees; D055549:Volatile Organic Compounds / Q000302:isolation & purification / Q000378:metabolism*
chemical_list: D013729:Terpenes; D055549:Volatile Organic Compounds