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Classic research on elevational gradients in plant–herbivore interactions holds that insect herbivore pressure is stronger under warmer, less seasonal climates characteristic of low elevations, and that this in turn selects for increased defence in low- (relative to high-) elevation plants. However, recent work has questioned this paradigm, arguing that it overly simplifies the ecological complexity in which plant–insect herbivore interactions are embedded along elevational gradients. Numerous biotic and abiotic factors vary with elevation, and their simultaneous influences are the focus of current work on elevational gradients in insect herbivory and plant defences. The present review 1) synthesizes current knowledge on elevational gradients in plant–insect herbivore interactions; 2) critically analyses research gaps and highlights recent advances that contribute to filling these gaps; and 3) outlines new research opportunities to uncover underlying mechanisms and build towards a unified theory on elevational gradients. We conclude that the next generation of studies should embrace community complexity – including multi-trophic dynamics and the multivariate nature of plant defence – and to do so by combining observational data, manipulative experiments and emerging analytical tools.

There is abundant evidence that leaf mechanical traits deter feeding by insect herbivores, but little is known about which particular traits contribute to defence across feeding guilds. We investigated the contribution of multiple mechanical traits from shear, punch and tear tests to herbivore deterrence across feeding guilds. Visible damage from miners and external chewers was measured and sucker feeding density estimated in mature leaves of 20 species of forest shrubs and small trees. Cafeteria trials were undertaken using a generalist chewer (larvae of Epiphyas postvittana, Lepidoptera). Damage was compared with leaf mechanical traits and associated nutrient and chemical defence traits. Damage by external chewers in the field and by E. postvittana correlated negatively with mechanical traits. Hierarchical partitioning analysis indicated that the strongest independent contribution to chewing damage was by the material trait of specific work to shear, with 68 % of total variance explained by the combination of specific work to shear (alone explaining 54 %) and tannin activity in a regression model. Mining damage did not correlate with mechanical traits, probably because miners can avoid tissues that generate high strength and toughness in mature leaves. Mechanical traits correlated more strongly with chewing damage in the field than chemical defences (total phenolics and tannin activity) and nutrients (nitrogen and water), but nutrients correlated strongly with diet selection in the cafeteria trial. Surprisingly, sucker feeding density correlated positively with mechanical traits and negatively with nutrients. Mechanical traits of mature leaves influenced insect feeding guilds differentially, reflecting differences in life history and feeding modes. For external chewers, energy (work) to fracture in shearing tests, at both structural and material levels, was strongly predictive of damage. Knowing which leaf mechanical traits influence insect feeding, and in which guilds, is important to our wider understanding of plant-herbivore interactions.

Acute and precise signal perception and transduction are essential for plant defense against insects. Insect elicitors-that is, the biologically active molecules from insects' oral secretion (which contains regurgitant and saliva), frass, ovipositional fluids, and the endosymbionts-are recognized by plants and subsequently induce a local or systematic defense response. On the other hand, insects secrete various types of effectors to interfere with plant defense at multiple levels for better adaptation. Jasmonate is a main regulator involved in plant defense against insects and integrates with multiple pathways to make up the intricate defense network. Jasmonate signaling is strictly regulated in plants to avoid the hypersensitive defense response and seems to be vulnerable to assault by insect effectors at the same time. Here, we summarize recently identified elicitors, effectors, and their target proteins in plants and discuss their underlying molecular mechanisms.

Increasing global food production is threatened by harsh environmental conditions along with biotic stresses, requiring massive new research into integrated stress resistance in plants. Stomata play a pivotal role in response to many biotic and abiotic stresses, but their orchestrated interactions at the molecular, physiological, and biochemical levels were less investigated. Here, we reviewed the influence of drought, pathogen, and insect herbivory on stomata to provide a comprehensive overview in the context of stomatal regulation. We also summarized the molecular mechanisms of stomatal response triggered by these stresses. To further investigate the effect of stomata–herbivore interaction at a transcriptional level, integrated transcriptome studies from different plant species attacked by different pests revealed evidence of the crosstalk between abiotic and biotic stress. Comprehensive understanding of the involvement of stomata in some plant–herbivore interactions may be an essential step towards herbivores’ manipulation of plants, which provides insights for the development of integrated pest management strategies. Moreover, we proposed that stomata can function as important modulators of plant response to stress combination, representing an exciting frontier of plant science with a broad and precise view of plant biotic interactions.

Abstract Various studies have addressed the impact of microbial inoculants on the composition of the resident microbiome. How microbial inoculants impact plant metabolism and interact with the resident rhizobiota under herbivory stress remains elusive. Here, we investigated the impact of two bacterial and two fungal inoculants, inoculated as single species and as a synthetic community, on the rhizosphere microbiome and volatilome of tomato plants (Solanum lycopersicum) comparing nonstress conditions to exposed to leaf herbivory by Spodoptera exigua. Based on amplicon sequencing analysis, rhizobacterial community composition was significantly affected by all four inoculants and the magnitude of this effect was dependent on herbivory stress. Fungal community composition was altered by the microbial inoculants but independent of herbivory stress. The rhizosphere volatilome was impacted by the microbial inoculation and differences between treatments were evened under herbivory stress. Each microbial inoculant caused unique changes in the volatilome of stressed plants but also shared similar responses, in particular the enhanced production of dimethyl disulfide and benzothiazole. In conclusion, the introduction of microbial inoculants in the tomato rhizosphere caused unique as well as common changes in the rhizosphere microbiome and volatilome, but these changes were minor compared to the microbiome changes induced by herbivory stress. Changes in rhizospheric bacterial and fungal communities and volatilome of tomato plants under microbial inoculation of beneficial microbes as single species and as a synthetic community under leaf herbivory stress conditions.

Biodiversity effects on ecosystem functioning can be partitioned into complementarity effects, driven by many species, and selection effects, driven by few. Selection effects occur through interspecific abundance shifts (dominance) and intraspecific shifts in functioning. Complementarity and selection effects are often calculated for biomass, but very rarely for secondary productivity, that is, energy transfer to higher trophic levels. We calculated diversity effects for three functions: aboveground biomass, insect herbivory and pathogen infection, the latter two as proxies for energy transfer to higher trophic levels, in a grassland experiment (PaNDiv) manipulating species richness, functional composition, nitrogen enrichment, and fungicide treatment. Complementarity effects were, on average, positive and selection effects negative for biomass production and pathogen infection and multiple species contributed to diversity effects in mixtures. Diversity effects were, on average, less pronounced for herbivory. Diversity effects for the three functions were not correlated, because different species drove the different effects. Benefits (and costs) from growing in diverse communities, be it reduced herbivore or pathogen damage or increased productivity either due to abundance increases or increased productivity per area were distributed across different plant species, leading to highly variable contributions of single species to effects of diversity on different functions. These results show that different underlying ecological mechanisms can result in similar overall diversity effects across functions.

In addition to direct tissue consumption, herbivory may affect other important plant processes. Here, we evaluated the effects of short-time leaf feeding by Spodoptera exigua larvae on the photosynthetic efficiency of tomato plants, using chlorophyll a fluorescence imaging analysis. After 15 min of feeding, the light used for photochemistry at photosystem II (PSII) (ΦPSII), and the regulated heat loss at PSII (ΦNPQ) decreased locally at the feeding zones, accompanied by increased non-regulated energy losses (ΦNO) that indicated increased singlet oxygen (1O2) formation. In contrast, in zones neighboring the feeding zones and in the rest of the leaf, ΦPSII increased due to a decreased ΦNPQ. This suggests that leaf areas not directly affected by herbivory compensate for the photosynthetic losses by increasing the fraction of open PSII reaction centers (qp) and the efficiency of these centers (Fv’/Fm’), because of decreased non-photochemical quenching (NPQ). This compensatory reaction mechanism may be signaled by singlet oxygen formed at the feeding zone. PSII functionality at the feeding zones began to balance with the rest of the leaf 3 h after feeding, in parallel with decreased compensatory responses. Thus, 3 h after feeding, PSII efficiency at the whole-leaf level was the same as before feeding, indicating that the plant managed to overcome the feeding effects with no or minor photosynthetic costs.

Plant responses to insect eggs can result in intensified defences against hatching larvae. In annual plants, this egg-mediated effect is known to be associated with changes in leaf phenylpropanoid levels. However, little is known about how trees—long-living, perennial plants—improve their egg-mediated, anti-herbivore defences. The role of phytohormones and the phenylpropanoid pathway in egg-primed anti-herbivore defences of a tree species has until now been left unexplored. Using targeted and untargeted metabolome analyses we studied how the phenylpropanoid pathway of Ulmus minor responds to egg-laying by the elm leaf beetle and subsequent larval feeding. We found that when compared to untreated leaves, kaempferol and quercetin concentrations increased in feeding-damaged leaves with prior egg deposition, but not in feeding-damaged leaves without eggs. PCR analyses revealed that prior insect egg deposition intensified feeding-induced expression of phenylalanine ammonia lyase (PAL), encoding the gateway enzyme of the phenylpropanoid pathway. Salicylic acid (SA) concentrations were higher in egg-treated, feeding-damaged leaves than in egg-free, feeding-damaged leaves, but SA levels did not increase in response to egg deposition alone—in contrast to observations made of Arabidopsis thaliana. Our results indicate that prior egg deposition induces a SA-mediated response in elms to feeding damage. Furthermore, egg deposition boosts phenylpropanoid biosynthesis in subsequently feeding-damaged leaves by enhanced PAL expression, which results in the accumulation of phenylpropanoid derivatives. As such, the elm tree shows similar, yet distinct, responses to insect eggs and larval feeding as the annual model plant A. thaliana.

The properties of the human mind are responsible for a number of biases that affect the quality of scientific research. However, scientists working in the fields of ecology and environmental science rarely take these biases into account. We conducted a meta-analysis of data extracted from 125 publications comparing woody plant damage by defoliating insects in different environments in order to understand the extent to which our knowledge on spatial patterns in herbivory is affected by various biases. We asked which research methods are most prone to biases and whether these biases lead to overestimation of the effects under study. The effect sizes (ESs) decreased with increases in the numbers of plant species involved in the study, with 61% lower ESs for herbivory estimated on all plants growing in study plots compared to herbivory on selected species. ESs also depended on the leaf sampling procedure: when all leaves from a tree or branch were sampled for measurements of herbivory or when random or systematic selection protocols were applied, ESs were 74% smaller than in cases of more subjective haphazard selection. In addition, ESs were 97% and 135% greater when the person conducting sampling and measuring was aware of the research hypothesis or sample origin, when compared with situations when the observer was blinded to these factors. The impacts of cognitive biases on the study outcomes significantly decreased with the increase in publication year; however, this pattern emerged mostly due to high-ranked journals and was non-significant for other journals. Using the studies of spatial patterns in herbivory as an example, we showed that our ecological and environmental knowledge is considerably biased due to an unconscious tendency of researchers to find support for their hypotheses and expectations, which generally leads to overestimation of the effects under study. Cognitive biases can be avoided by using different methods, such as applying randomization procedures in sampling and blinding of research hypotheses and sample origins. These measures should be seen as obligatory; otherwise, accumulation of the biased results in primary studies may ultimately lead to false general conclusions in subsequent research synthesis.

The relationship between plant nutrient content and insect herbivore populations and community structure has long interested ecologists. Insect herbivores require multiple nutrients, but ecologists have focused mostly on nitrogen (an estimate of plant protein content), and more recently phosphorus (P); other nutrients have received little attention. Here we document nutrient variation in grass and forb samples from grassland habitats in central Nebraska using an elemental approach; in total we measured foliar concentrations of 12 elements (N and P, plus S, B, Ca, Mg, Na, K, Zn, Fe, Mn, and Cu). We detected significant variability among sites for N, P, Mg, Na, K, and Cu. We next used a model selection approach to explore how this nutritional variation and plant biomass correlate with grasshopper densities (collectively and at the feeding-guild level), and principal component analysis to explore nutrient correlations with grasshopper community species composition. When all grasshoppers were pooled, densities varied among sites, but only P was associated with abundance of the elements shown to vary between sites. Different responses occurred at the feeding-guild level. For grass specialists, densities were associated with N, plus P, Mg, and Na. For forb specialists, N and P were often associated with density, but associations with Na and K were also observed. Finally, mixed-feeder abundance was strongly associated with biomass, and to a lesser extent P, Mg, Na, and Cu. At the community level, B, Ca, Zn, and Cu, plus biomass, explained >30% of species composition variation. Our results confirm the positive association of N and P with insect herbivore populations, while suggesting a potential role for Mg, Na, and K. They also demonstrate the importance of exploring effects at the feeding-guild level. We hope our data motivate ecologists to think beyond N and P when considering plant nutrient effects on insect herbivores, and make a call for studies to examine functional responses of insect herbivores to dietary manipulation of Mg, Na, and K. Finally, our results demonstrate correlations between variation in nutrients and species assemblages, but factors not linked to plant nutrient quality or biomass likely explain most of the observed variation.