title: Environmental influences on growth and defence responses of the invasive shrub, Lonicera maackii, to simulated and real herbivory in the juvenile stage.
abstract: OBJECTIVE
  Tolerance and defence against herbivory are among the many mechanisms attributed to the success of invasive plants in their novel ranges. Because tolerance and defence against herbivory differ with the ontogeny of a plant, the effects of herbivore damage on plant fitness vary with ontogenetic stage and are compounded throughout a plant's lifetime. Environmental stresses such as light and nutrient limitations can further influence the response of the plant. Much is known about the response of plants in the seedling and reproductive adult stages, but less attention has been given to the pre-reproductive juvenile stage.
  
  
  METHODS
  Juvenile plants of the North American invasive Lonicera maackii were exposed to simulated herbivory under high and low light and nitrogen availability and growth, allocation patterns and foliar defensive chemistry were measured. In a second experiment, complete nutrient availability and damage type (generalist caterpillar or simulated) were manipulated.
  
  
  RESULTS
  Juvenile plants receiving 50 % defoliation had lower total biomass and a higher root^:^shoot ratio than controls for all treatment combinations except low nitrogen/low light. Low light and defoliation increased root^:^shoot ratio. Light, fertilization and defoliation had little impact on foliar defensive chemistry. In the second experiment, there was a reduction in total biomass when caterpillar damage was applied. The root^:^shoot ratio increased under low soil fertility and was not affected by defoliation. Stem-diameter growth rates and specific leaf area did not vary by damage type or fertilization. Foliar protein increased through time, and more strongly in defoliated plants than in controls, while peroxidase activity and total flavonoids decreased with time. Overall, resource limitations were more influential than damage in the growth of juvenile L. maackii plants.
  
  
  CONCLUSIONS
  The findings illustrate that even when resources are limited, the tolerance and defence against herbivory of a woody invasive plant in the juvenile stage may contribute to the establishment and persistence of some species in a variety of habitats.
authors.affiliation: Department of Biological Sciences and Environmental Sciences PhD Program, Wright State University, 3640 Colonel Glenn Highway, Dayton, OH 45435, USA. dmlieurance@ufl.edu
authors.forename: Deah
authors.initials: D
authors.lastname: Lieurance
authors.forename: Don
authors.initials: D
authors.lastname: Cipollini
mesh_terms: D000818:Animals; D005419:Flavonoids / Q000378:metabolism; D060434:Herbivory; D008027:Light; D031239:Lonicera / Q000254:growth & development* / Q000378:metabolism; D009584:Nitrogen / Q000378:metabolism*; D009195:Peroxidase / Q000378:metabolism; D018515:Plant Leaves / Q000378:metabolism; D010940:Plant Proteins / Q000378:metabolism
chemical_list: D005419:Flavonoids; D010940:Plant Proteins; D009195:Peroxidase; D009584:Nitrogen
keywords: Herbivory; Lonicera maackii; ontogeny; relative growth rate; root to shoot ratio; tolerance