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The resource‐availability hypothesis (RAH) and the intraspecific RAH (RAH intra ), posit that resources, (i.e. nutrients) control plant antiherbivore defenses. Both hypotheses predict that in low‐resource environments, plant growth is slow, and constitutive defense is high. In high‐resource environments, however, the RAH predicts that plant growth is fast, and constitutive defense is low, whereas the RAH intra predicts that increased resources attract more herbivores, and this intensified grazing pressure leads to high constitutive defense. Salt marshes are nutrient‐limited ecosystems threatened by eutrophication and chronic herbivory, yet we know little about how these stressors shape saltmarsh plant antiherbivore defenses, which influence trophic interactions and ecosystem resilience. We manipulated resource availability via nutrient addition and herbivory via the marsh periwinkle Littoraria irrorata , on the saltmarsh foundation species Spartina alterniflora , in mesocosms. Because plant age can also influence trait variation, we measured traits in both original and clonally‐grown new stems. Feeding assays then evaluated how treatments and plant age affected subsequent Littoraria consumption of Spartina . Nutrient addition stimulated growth, while decreasing defensive traits (e.g. fiber and silica content), following the RAH. Herbivory enhanced belowground production and increased stem diameter, yet did not induce defensive traits, contrary to our expectations. Herbivory plus nutrients increased Spartina biomass and reduced phenolics, a defensive trait, further supporting the RAH. Regardless of treatment, clonally‐grown new stems had greater variation in measured traits. Despite altered traits, however, treatments and plant age did not affect Littoraria consumption. Our results support the RAH and part of the RAH intra and suggest: 1) nutrient availability is a primary driver of plant trait change and 2) plant age controls the magnitude of trait variation in Spartina . Further, our findings indicate that eutrophic conditions may not always increase top‒down control by herbivores, and in some instances can enhance saltmarsh resilience against sea‐level rise via stimulated Spartina biomass production.
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This content will become publicly available on July 17, 2026
Three decades of experimental nitrogen fertilization increases the frequency of a defensive plant morph
Resources, such as nitrogen, are widely hypothesized to underlie the expression and evolution of plant defenses to herbivory. However, resource availability can affect selection on plant defense traits in contrasting ways: resource availability can 1) weaken selection on defense traits by reducing the costs of herbivory, or 2) strengthen selection on defense traits by increasing herbivore pressure. Previous studies have compared herbivore resistance in populations across natural resource gradients to infer how resource availability affects the microevolution of plant defenses. However, because these studies do not manipulate resource availability, they are unable to directly test the effects of resources of plant defense trait evolution. We used a three‐decade‐long nitrogen fertilization field experiment to test how nitrogen availability affects the evolution of an architectural plant defense trait: stem nodding inSolidago altissima. Stem nodding is a genetic dimorphism that helps plants to evade apex‐galling herbivores. By comparing the frequency of defensive nodding versus erect morphs in experimentally fertilized or unfertilized plots 27, 32 and 33 years post‐treatment initiation, we assessed how nitrogen addition affects the evolution of this defense trait. We found that the defensive nodding morph was 3–6 times more common in plots that evolved under nitrogen fertilization compared to those that evolved in unfertilized control plots. This study provides empirical evidence for resource availability driving plant defense evolution and demonstrates that this evolution can occur on time‐scales conducive to study at many long‐term nutrient fertilization experiments.
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- Award ID(s):
- 2224712
- PAR ID:
- 10635568
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Oikos
- ISSN:
- 0030-1299
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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