We present a framework to explain how prey stress responses to predation can resolve context dependency in ecosystem properties and functions such as food chain length, secondary production, elemental stoichiometry, and cycling. We first describe the major nonspecific physiological stress mechanisms and their ecologically relevant consequences. We next synthesize the evidence for prey physiological responses to predation risk and demonstrate that they are similar across taxa and fit well within the general stress paradigm. We then illustrate the utility of our idea by applying our understanding of the ecological consequences of stress to explain how herbivore‐prey physiological antipredator responses affect ecosystem dynamics. We hypothesize that stressed herbivores should forage on plant species with higher digestible carbohydrates than should unstressed herbivores to meet heightened energy demands. Increased consumption of carbohydrate‐rich plants should reduce their relative abundance in the community, hence altering the quantity and quality of plant litter entering the detrital pool. We further hypothesize that stress should change the elemental composition and energy content of prey excreta, egesta, and carcasses that enter the detrital pool. Finally, prey stress should lower energy and nutrient conversion efficiency and hence the transfer of materials and energy up the food chain, which should, in turn, weaken the association between ecosystem productivity and food chain length.
Predators are predominantly valued for their ability to control prey, as indicators of high levels of biodiversity and as tourism attractions. This view, however, is incomplete because it does not acknowledge that predators may play a significant role in the delivery of critical life-support services such as ecosystem nutrient cycling. New research is beginning to show that predator effects on nutrient cycling are ubiquitous. These effects emerge from direct nutrient excretion, egestion or translocation within and across ecosystem boundaries after prey consumption, and from indirect effects mediated by predator interactions with prey. Depending on their behavioural ecology, predators can create heterogeneous or homogeneous nutrient distributions across natural landscapes. Because predator species are disproportionately vulnerable to elimination from ecosystems, we stand to lose much more from their disappearance than their simple charismatic attractiveness.
The process of nutrient transfer through an ecosystem is an important determinant of production, food-chain length, and species diversity. The general view is that the rate and efficiency of nutrient transfer up the food chain is constrained by herbivore-specific capacity to secure N-rich compounds for survival and production. Using feeding trials with artificial food, we show, however, that physiological stressresponse of grasshopper herbivores to spider predation risk alters the nature of the nutrient constraint. Grasshoppers facing predation risk had higher metabolic rates than control grasshoppers. Elevated metabolism accordingly increased requirements for dietary digestible carbohydrate-C to fuel-heightened energy demands. Moreover, digestible carbohydrate-C comprises a small fraction of total plant tissue-C content, so nutrient transfer between plants and herbivores accordingly becomes more constrained by digestible plant C than by total plant C:N. This shift in herbivore diet to meet the altered nutrient requirement increased herbivore body C:N content, the C:N content of the plant community from which grasshoppers select their diet, and grasshopper fecal C:N content. Chronic predation risk thus alters the quality of animal and plant tissue that eventually enters the detrital pool to become decomposed. Our results demonstrate that herbivore physiology causes C:N requirements and nutrient intake to become flexible, thereby providing a mechanism to explain context dependence in the nature of trophic control over nutrient transfer in ecosystems.ecological stoichiometry | metabolism | nutrient balance | physiological stress | predator-prey interaction
Aboveground consumers are believed to affect ecosystem functioning by regulating the quantity and quality of plant litter entering the soil. We uncovered a pathway whereby terrestrial predators regulate ecosystem processes via indirect control over soil community function. Grasshopper herbivores stressed by spider predators have a higher body carbon-to-nitrogen ratio than do grasshoppers raised without spiders. This change in elemental content does not slow grasshopper decomposition but perturbs belowground community function, decelerating the subsequent decomposition of plant litter. This legacy effect of predation on soil community function appears to be regulated by the amount of herbivore protein entering the soil.
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