Kessler et al. 2010 Changing Pollinators as a Means of Escaping Herbivores (1)
Summary
Overview
Animal-pollinated plants face the challenge of attracting pollinators without attracting herbivores.
This dilemma is heightened when larval-stage herbivores also serve as important pollinators.
Nicotiana attenuata
Nicotiana attenuata is a night-flowering tobacco that typically opens flowers at night to attract nocturnal hawkmoth pollinators (e.g., Manduca quinquemaculata, Manduca sexta).
When infested by hawkmoth larvae, these plants change their floral characteristics.
Herbivore Impact on Flower Phenology
Upon larval attack, N. attenuata produces morning-opening flowers (MoFs) instead of night-opening flowers (NoFs).
MoFs exhibit reduced emissions of floral volatiles (like benzyl acetone) and attract diurnal pollinators, specifically hummingbirds.
This shift in flowering time is regulated by oral secretions from the feeding larvae and requires jasmonate (JA) signaling.
Experimental Observations
Field Studies
During a Manduca quinquemaculata outbreak, plants shifted from producing primarily NoFs to MoFs in response to herbivore attack.
Experiments showed that infested plants produced significantly more MoFs under herbivore pressure compared to uninfested plants.
Flower Characteristics
MoFs have:
Lower benzyl acetone emissions.
Reduced nectar concentrations.
Smaller corolla diameter compared to NoFs.
The shift from NoFs to MoFs affects the community of pollinators visiting the plants, facilitating greater pollination by hummingbirds.
Pollinator Dynamics
Hummingbird Visits
Hummingbirds preferentially visit MoFs over NoFs, likely due to nectar availability that is undisturbed by nocturnal pollinators.
Observational data highlighted that visits to MoFs increased overall pollination rates of infested plants.
Fitness Consequences
Infested plants benefited from enhanced outcrossing rates despite emitting lower floral volatiles overall.
Hummingbirds’ preferences contributed to a greater capsule production from MoFs compared to NoFs.
Mechanisms of Change
Signaling Pathways
Feeding by Manduca larvae activates a MAP kinase cascade, leading to increased jasmonic acid production and resulting in the herbivory-induced production of MoFs.
Genetic experiments confirmed that JA signaling is essential for this change in flower phenology.
Conclusion
The shift from MoFs to NoFs serves as a defensive strategy against herbivore attack while optimizing pollination services from specific pollinators such as hummingbirds.
This herbivory-elicited change in flower phenology accentuates the complex interactions between plants, herbivores, and pollinators, representing an adaptation for reducing herbivore loads while maximizing reproductive success.