title: Age-dependent induction of immunity and subsequent survival costs in males and females of a temperate damselfly.
abstract: BACKGROUND
  To understand variation in resistance to parasites within host populations, researchers have examined conditions under which immunity is induced and/or is costly. Both host sex and age have been found to influence immune expression and subsequently are likely factors influencing the costs of resistance. The purpose of this study was to examine immune expression and associated survival costs for two age groups (newly emerged and sexually mature individuals) of the damselfly, Enallagma boreale Selys. Survival was assessed for experimentally challenged and control damselflies, housed initially at 22 degrees C and then subjected to low temperatures (15 degrees C) associated with reduced foraging activity and food deprivation. Experimental conditions emulated natural local variation in bouts of good weather followed by inclement weather (successions of days with hourly mean temperatures around 15 degrees C and/or rainy weather).
  
  
  RESULTS
  At least one of three immune traits was induced to higher levels for both newly emerged and mature E. boreale challenged by Lippopolysaccharide (LPS) relative to saline-injected controls, when housed at 22 degrees C. The immune traits assayed included haemocyte concentration, Phenoloxidase activity and antibacterial activity and their induction varied among ages and between males and females. For matures, those injected with LPS had lowered survivorship compared to saline-injected controls that were housed initially at 22 degrees C and subsequently at 15 degrees C. Newly emerged LPS-injected damselflies did not show reduced survivorship relative to newly-emerged controls, despite showing immune induction.
  
  
  CONCLUSIONS
  Reduced longevity following induction of immunity was observed for reproductively mature damselflies, but not for newly emerged damselflies. Costs of resistance depend only partly on the immune trait induced and more on the age (but not sex) of the host. In four years, we often observed bouts of inclement weather following good days and these bouts occurred primarily during the emergence periods, but also during the flight periods, of E. boreale. The duration of these bouts appear sufficient to compromise survival of mature damselflies that responded immunologically to LPS challenge. We further suggest the environmental conditions likely experienced by different ages of damselflies, following resistance expression, has influenced optimal immune investment by individuals in different age classes and the likelihood of detecting costs of resistance.
authors.affiliation: Department of Ecology & Evolutionary Biology, 25 Willcocks Street, University of Toronto, Toronto, Ontario, M5S 3B2, Canada. trobb@connect.carleton.ca
authors.forename: Tonia
authors.initials: T
authors.lastname: Robb
authors.forename: Mark R
authors.initials: MR
authors.lastname: Forbes
mesh_terms: D000367:Age Factors; D000818:Animals; D000900:Anti-Bacterial Agents / Q000378:metabolism; D005260:Female; D006434:Hemocytes / Q000378:metabolism; D007113:Immunity, Innate / Q000502:physiology; D007313:Insecta / Q000276:immunology* / Q000378:metabolism / Q000502:physiology; D008070:Lipopolysaccharides / Q000494:pharmacology; D008297:Male; D014442:Monophenol Monooxygenase / Q000378:metabolism; D012737:Sex Factors; D013696:Temperature
chemical_list: D000900:Anti-Bacterial Agents; D008070:Lipopolysaccharides; D014442:Monophenol Monooxygenase