The carbon–nutrient balance hypothesis: its rise and fall

The idea that the concentration of secondary metabolites in plant tissues is controlled by the availability of carbon and nitrogen in the environment has been termed the carbon–nutrient balance hypothesis (CNB). This hypothesis has been invoked both for prediction and for post hoc explanation of the results of hundreds of studies. Although it successfully predicts outcomes in some cases, it fails to such an extent that it cannot any longer be considered useful as a predictive tool. As information from studies has accumulated, many attempts have been made to save CNB, but these have been largely unsuccessful and have managed only to limit its utility. The failure of CNB is rooted in assumptions that are now known to be incorrect and it is time to abandon CNB because continued use of the hypothesis is now hindering understanding of plant–consumer interactions. In its place we propose development of theory with a firm evolutionary basis that is mechanistically sophisticated in terms of plant and herbivore physiology and genetics.     

The function of the wild carrot's dark central floret: attract,guide or deter?

Although Charles Darwin wrote about flower polymorphism in the wild carrot, Daucus carota, the function of the conspicuous central dark floret is still unclear. We used video recordings to evaluate the functional significance of the dark central floret as a short‐distance signal for insects landing on the umbels and analyzed the location of landing points, landing orientation and visit duration. These parameters, as well as insect attraction did not differ on umbels with and without a dark central floret. Hence, we found no evidence for the role of the dark central floret in the pollination of D. carota. Umbels with a dark central floret, however, were parasitized significantly less often by the gall midge Kiefferia pericarpiicola than umbels without a dark central floret. We propose that the dark central floret may play a role in reducing parasite infestation by mimicking an already present gall or deterring oviposition of the gall midge by other means.     

Effects of Nitrogen Deposition on Insect Herbivory: Implications for Community and Ecosystem Processes

The deposition of anthropogenically fixed nitrogen (N) from the atmosphere onto land and plant surfaces has strong influences on terrestrial ecosystem processes. Although recent research has expanded our understanding of how N deposition affects ecosystems directly, less attention has been directed toward the investigation of how N deposition may affect ecosystems indirectly by modifying interactions among organisms. Empirical evidence suggests that there are several mechanisms by which N deposition may affect interactions between plants and insect herbivores. The most likely mechanisms are deposition-induced shifts in the quality and availability of host plant tissues. We discuss the effects of N deposition on host plant chemistry, production, and phenology, and we review the evidence for the effects of N deposition on insect herbivores at the individual, population, and community levels. In general, N deposition has positive effects on individual insect performance, probably due to deposition-induced improvements in host plant chemistry. These improvements include increased N and decreased carbon-based defensive compound concentrations. The evidence to date suggests that N deposition may also have a positive effect on insect populations. These effects may have considerable ecological, as well as economic consequences if the rates of herbivory on economically important timber species continue to increase. Deposition-induced changes in plant–herbivore relationships may affect community and ecosystem processes. However, we predict that the larger-scale consequences of interactions between N deposition and herbivory will vary based on site-specific factors. In addition, interactions between N deposition and other global-scale changes may lead to nonadditive effects on patterns of herbivory.     

Seedling–herbivore interactions: insights into plant defence and regeneration patterns

Background

Herbivores have the power to shape plant evolutionary trajectories, influence the structure and function of vegetation, devastate entire crops, or halt the spread of invasive weeds, and as a consequence, research into plant–herbivore interactions is pivotal to our understanding of plant ecology and evolution. However, the causes and consequences of seedling herbivory have received remarkably little attention, despite the fact that plants tend to be most susceptible to herbivory during establishment, and this damage can alter community composition and structure.

Scope

In this Viewpoint article we review why herbivory during early plant ontogeny is important and in so doing introduce an Annals of Botany Special Issue that draws together the latest work on the topic. In a synthesis of the existing literature and a collection of new studies, we examine several linked issues. These include the development and expression of seedling defences and patterns of selection by herbivores, and how seedling selection affects plant establishment and community structure. We then examine how disruption of the seedling–herbivore interaction might affect normal patterns of plant community establishment and discuss how an understanding of patterns of seedling herbivory can aid our attempts to restore semi-natural vegetation. We finish by outlining a number of areas where more research is required. These include a need for a deeper consideration of how endogenous and exogenous factors determine investment in seedling defence, particularly for the very youngest plants, and a better understanding of the phylogenetic and biogeographical patterns of seedling defence. There is also much still be to be done on the mechanisms of seedling selection by herbivores, particularly with respect to the possible involvement of volatile cues. These inter-related issues together inform our understanding of how seedling herbivory affects plant regeneration at a time when anthropogenic change is likely to disrupt this long-established, but all-too-often ignored interaction.     

No post-Cretaceous ecosystem depression in European forests? Rich insect-feeding damage on diverse middle Palaeocene plants,Menat, France

Insect herbivores are considered vulnerable to extinctions of their plant hosts. Previous studies of insect-damaged fossil leaves in the US Western Interior showed major plant and insect herbivore extinction at the Cretaceous–Palaeogene (K–T) boundary. Further, the regional plant–insect system remained depressed or ecologically unbalanced throughout the Palaeocene. Whereas Cretaceous floras had high plant and insect-feeding diversity, all Palaeocene assemblages to date had low richness of plants, insect feeding or both. Here, we use leaf fossils from the middle Palaeocene Menat site, France, which has the oldest well-preserved leaf assemblage from the Palaeocene of Europe, to test the generality of the observed Palaeocene US pattern. Surprisingly, Menat combines high floral diversity with high insect activity, making it the first observation of a ‘healthy’ Palaeocene plant–insect system. Furthermore, rich and abundant leaf mines across plant species indicate well-developed host specialization. The diversity and complexity of plant–insect interactions at Menat suggest that the net effects of the K–T extinction were less at this greater distance from the Chicxulub, Mexico, impact site. Along with the available data from other regions, our results show that the end-Cretaceous event did not cause a uniform, long-lasting depression of global terrestrial ecosystems. Rather, it gave rise to varying regional patterns of ecological collapse and recovery that appear to have been strongly influenced by distance from the Chicxulub structure.     

Choice,performance and heritability of performance of specialist and generalist insect herbivores towards cacalol and seneciphylline,two allelochemicals of Adenostyles alpina (Asteraceae)

We compared the effects of a sesquiterpene (ST, cacalol) and a pyrrolizidine alkaloid (PA, seneciphylline), both occurring in Adenostyles alliariae, on food choice and performance of specialist and generalist insect herbivores which are all known to feed or live on A. alliariae. In choice experiments we investigated whether the compounds were preferred, deterrent or had no effect. All specialist species Aglaostigma discolor (Hymenoptera, Tenthredinidae), Oreina cacaliae (Coleoptera, Chrysomelidae) and O. speciosissima avoided feeding when confronted with the combination of compounds. Only larvae of A. discolor avoided the single ST treatment as well. Larvae of the generalist species Callimorpha dominula (Lepidoptera, Arctiidae), Cylindrotoma distinctissima (Diptera, Tipulidae) and Miramella alpina (Caelifera, Acrididae) generally avoided feeding from PA, ST and PAST treatments. The only exception were caterpillars of C. dominula which were indiscriminate towards PA when naive, and preferred to feed on the PA treatment when they had experienced the compound before. Performance, measured as the growth of larvae on the different treatments in a no choice situation over a period of 10–17 days, was not different between treatments in the specialist leaf beetles O. cacaliae and O. speciosissima. Their avoidance of the combination treatment in the choice experiments had no obvious effect on growth when forced to feed from the treatment. In the generalist C. dominula only the high concentration combination treatment (PAST) reduced growth of the larvae due to decreased consumption. In C. distinctissima we found reduced growth in all treatments except one (PA3%). Poor growth performance in C. distinctissima was due to postingestive physiological effects of all treatments and additionally to consumption reduction in high‐dose ST treatments. Genetic variability (broad sense heritability) of growth performance metabolism varied in accordance with the specialization degree of the species. O. cacaliae, the most specialized species, had no significant heritability; O. speciosissima, the less specialized specialist, had a heritability of 0.46; C. dominula, the PA adapted generalist species, had a heritability of 0.64; C. distinctissima, the generalist with no apparent adaptations, had a heritability of 0.84.