The defensive role of volatile emission and extrafloral nectar secretion for lima bean in nature.

Lima bean (Phaseolus lunatus) features two indirect anti-herbivore defenses--emission of volatile organic compounds (VOCs) and secretion of extrafloral nectar (EFN)--which are both inducible upon herbivore damage. In a previous field study, Lima bean benefited from the simultaneous induction of the two defenses, yet it remained unclear whether both had contributed to plant protection. Our experimental approach aimed at studying the defensive role of both indirect defenses simultaneously. Tendrils were sprayed with jasmonic acid (JA) to induce both defenses, and performance was compared to that of others that were treated with a synthetic blend of either EFN or VOCs. Confirming earlier results, JA treatment and application of the VOC mixture induced EFN secretion in treated tendrils in quantitatively similar amounts. The composition of the applied synthetic blend of EFN was adjusted to match the concentration of EFN secreted from JA- and VOC-treated tendrils. Repeated application of either enhanced the performance of several fitness-relevant plant parameters such as growth rate and flower production. Tendrils treated with JA showed a similar trend, yet some fitness-related parameters responded less to this treatment. This suggests a minor importance of any putative JA-dependent direct defense traits or higher costs of JA-elicited responses as compared to VOCS and EFN, as otherwise JA-treated tendrils should have outperformed VOC- and EFN-treated tendrils. Moreover, the beneficial effect of applying synthetic EFN alone equaled or exceeded that of VOCs and JA. Ants were by far the dominant group among the arthropods that was attracted to JA-, VOC-, or EFN-treated tendrils. The results suggest that EFN plays a more important role as an indirect defense of lima bean than VOCs or any other JA-responsive trait.