Impact of the female and hermaphrodite forms of Opuntia robusta on the plant defence hypothesis

The optimal defence hypothesis predicts that increased plant defence capabilities, lower levels of damage, and lower investment in vegetative biomass will occur more frequently in sexual forms with higher resource-demanding tissue production and/or younger plant parts. We aimed to examine the effects of sexual form, cladode, and flower age on growth rate, herbivore damage, and 4-hydroxybenzoic acid (4-HBA), chlorogenic acid, and quercetin (QUE) concentrations in Opuntia robusta plants in central Mexico. Our findings demonstrated that hermaphrodite flowers showed faster growth and lesser damage than female flowers. The effect of cladode sexual forms on 4-HBA and QUE occurrence was consistent with the predictions of the optimal defence hypothesis. However, chlorogenic acid occurrences were not significantly affected by sexual forms. Old cladodes exhibited higher QUE and 4-HBA occurrences than young cladodes, and hermaphrodites exhibited higher 4-HBA concentrations than females. Resource allocation for reproduction and secondary metabolite production, and growth was higher and lower, respectively, in females, compared to hermaphrodites, indicating a trade-off between investment in reproduction, growth, and secondary metabolite production. Secondary metabolite concentrations in O. robusta plants were not negatively correlated with herbivore damage, and the two traits were not accurate predictors of plant reproductive output.

Florivory defence: are phenolic compounds distributed non-randomly within perianths?

Plants might allocate chemical defences unequally within attractive units of flowers including petals, sepals, and bracts because of variations in the probability of florivory. Based on optimal defence theory, which predicts that plants allocate higher chemical defences to tissues with higher probabilities of herbivore attack, we predicted that distal parts and sepals would have higher chemical defence allocations than proximal parts and petals. To test this prediction, we compared total phenolics and condensed tannins concentrations as well as presence of florivory within attractive units of ten angiosperm species. In agreement with the prediction, the overall results showed that the distal parts had higher total phenolics and condensed tannins than the proximal parts. On the other hand, contrary to the prediction, petals and sepals showed no tissue-specific variations. Florivory was more severe on the distal parts than the proximal parts, although statistical support for the variation was slightly weak, while the variations in presence of florivory between petals and sepals differed between the distal and proximal parts. These results may support the prediction of the optimal defence theory because distal parts of attractive units had higher presence of florivory and concentration of chemical defences.

Differential phenotypic and genetic expression of defence compounds in a plant–herbivore interaction along elevation

Elevation gradients impose large differences in abiotic and biotic conditions over short distances, in turn, likely driving differences in gene expression more than would genetic variation per se , as natural selection and drift are less likely to fix alleles at such a narrow spatial scale. As elevation increases, the pressure exerted on plants by herbivores and on arthropod herbivores by predators decreases, and organisms spanning the elevation gradient are thus expected to show lower levels of defence at high elevation. The alternative hypothesis, based on the optimal defence theory, is that defence allocation should be higher in low-resource habitats such as those at high elevation, due to higher costs associated with tissue replacement. In this study, we analyse variation with elevation in (i) defence compound content in the plant Lotus corniculatus and (ii) gene expression associated with defence against predators in the specific phytophagous moth, Zygaena filipendulae . Both species produce cyanogenic glycosides (CNglcs) such as lotaustralin and linamarin as defence mechanisms, with the moth, in addition, being able to sequester CNglcs from its host plant. Specifically, we tested the assumption that the defence-associated phenotype in plants and the gene expression in the insect herbivore should covary between low- and high-elevation environments. We found that L. corniculatus accumulated more CNglcs at high elevation, a result in agreement with the optimal defence theory. By contrast, we found that the levels of expression in the defence genes of Z. filipendulae larvae were not related to the CNglc content of their host plant. Overall, expression levels were not correlated with elevation either, with the exception of the UGT33A1 gene, which showed a marginally significant trend towards higher expression at high elevation when using a simple statistical framework. These results suggest that the defence phenotype of plants against herbivores, and subsequent herbivore sequestration machineries and de novo production, are based on a complex network of interactions.

Targeted predation of extrafloral nectaries by insects despite localized chemical defences

Extrafloral (EF) nectaries recruit carnivorous arthropods that protect plants from herbivory, but they can also be exploited by nectar thieves. We studied the opportunistic, targeted predation (and destruction) of EF nectaries by insects, and the localized chemical defences that plants presumably use to minimize this effect. In field and laboratory experiments, we identified insects that were possibly responsible for EF nectary predation in Vicia faba (fava bean) and determined the extent and accuracy of the feeding damage done to the EF nectaries by these insects. We also performed biochemical analyses of plant tissue samples in order to detect microscale distribution patterns of chemical defences in the area of the EF nectary. We observed selective, targeted feeding on EF nectaries by several insect species, including some that are otherwise not primarily herbivorous. Biochemical analyses revealed high concentrations of l -3,4-dihydroxyphenylalanine, a non-protein amino acid that is toxic to insects, near and within the EF nectaries. These results suggest that plants allocate defences to the protection of EF nectaries from predation, consistent with expectations of optimal defence theory, and that this may not be entirely effective, as insects limit their exposure to these defences by consuming only the secreting tissue of the nectary.