title: Temperature dependence of predation stress and the nutritional ecology of a generalist herbivore.
abstract: Prey at risk of predation may experience stress and respond physiologically by altering their metabolic rates. Theory predicts that such physiological changes should alter prey nutrient demands from N-rich to C-rich macronutrients and shift the balance between maintenance and growth/reproduction. Theory further suggests that for ectotherms, temperature stands to exacerbate this stress. Yet, the interactive effects of predation stress and temperature stress on diet, metabolism, and survival of ectotherms are not well known. This knowledge gap was addressed with a laboratory study in which wild juvenile grasshoppers were collected, assigned to one of three groups, and raised at three different temperatures. All grasshoppers had access to equal quantities of two diets composed of opposite carbohydrate : protein ratios. Half of the individuals in each temperature group were exposed to predation risk cues from spider predators, while the other half were kept in risk free conditions. Grasshoppers consumed more carbohydrates when exposed to predation risk, but consumption favored greater protein intake as temperature increased. Moreover, the difference in carbohydrate intake between risk cue and risk free treatments diminished as temperature increased. Furthermore, variability between individual consumption patterns both within and between treatments decreased markedly as temperature increased, suggesting that higher temperatures promote more consistent individual consumption behaviors. Grasshoppers grew faster and larger as temperature increased, which translated into higher survival rates at higher temperatures. Warmer grasshoppers also did not alter their metabolic rates in response to predation risk cues, in contrast to colder grasshoppers. Digestive efficiency increased with temperature as well -- further indicating that lower temperatures were much more stressful than higher temperatures for grasshoppers. The study shows that physiological responses of ectothermic herbivores to predation stress are highly plastic and temperature dependent, with higher temperatures promoting increased protein intake, growth, development, survival, and digestive efficiency relative to colder temperatures. These findings help to reconcile why dietary responses (proportion of protein vs. carbohydrate intake) to predation stress may vary among different prey taxa studied previously.
authors.affiliation: School of Forestry and Environmental Studies, Yale University, 370 Prospect Street, New Haven, Connecticut, 06511, USA.
authors.forename: Oswald J
authors.initials: OJ
authors.lastname: Schmitz
authors.affiliation: School of Forestry and Environmental Studies, Yale University, 370 Prospect Street, New Haven, Connecticut, 06511, USA.
authors.forename: Adam E
authors.initials: AE
authors.lastname: Rosenblatt
authors.affiliation: School of Forestry and Environmental Studies, Yale University, 370 Prospect Street, New Haven, Connecticut, 06511, USA.
authors.forename: Meredith
authors.initials: M
authors.lastname: Smylie
mesh_terms: D000818:Animals; D017753:Ecosystem; D004924:Escape Reaction; D006110:Grasshoppers / Q000502:physiology*; D060434:Herbivory; D011235:Predatory Behavior; D013112:Spiders / Q000502:physiology; D013312:Stress, Physiological; D013696:Temperature
keywords: 
  Melanoplus femurrubrum
  ; 
  Pisaurina mira
  ; carbohydrate; growth rate; metabolism; predation stress; protein; respirometry; thermal performance