Showing papers on "Plant tolerance to herbivory published in 2016"
TL;DR: Physical defense traits, plant vigor and herbivore-induced plant volatiles show considerable utility in pest control, along with mixed species crops, and emerging approaches for accelerating the identification of plant defensive traits and facilitating their deployment are highlighted to improve the future sustainability of crop protection.
Abstract: Interactions between plants and insect herbivores are important determinants of plant productivity in managed and natural vegetation. In response to attack, plants have evolved a range of defenses to reduce the threat of injury and loss of productivity. Crop losses from damage caused by arthropod pests can exceed 15% annually. Crop domestication and selection for improved yield and quality can alter the defensive capability of the crop, increasing reliance on artificial crop protection. Sustainable agriculture, however, depends on reduced chemical inputs. There is an urgent need, therefore, to identify plant defensive traits for crop improvement. Plant defense can be divided into resistance and tolerance strategies. Plant traits that confer herbivore resistance typically prevent or reduce herbivore damage through expression of traits that deter pests from settling, attaching to surfaces, feeding and reproducing, or that reduce palatability. Plant tolerance of herbivory involves expression of traits that limit the negative impact of herbivore damage on productivity and yield. Identifying the defensive traits expressed by plants to deter herbivores or limit herbivore damage, and understanding the underlying defense mechanisms, is crucial for crop scientists to exploit plant defensive traits in crop breeding. In this review, we assess the traits and mechanisms underpinning herbivore resistance and tolerance, and conclude that physical defense traits, plant vigor and herbivore-induced plant volatiles show considerable utility in pest control, along with mixed species crops. We highlight emerging approaches for accelerating the identification of plant defensive traits and facilitating their deployment to improve the future sustainability of crop protection.
239 citations
TL;DR: It is proposed that this phenomenon could play a key role in the suppression of herbivore populations in natural systems, and that increased nutrient heterogeneity within agricultural crops could contribute to the sustainable control of insect pests in agroecosystems.
Abstract: The performance and population dynamics of insect herbivores depend on the nutritive and defensive traits of their host plants. The literature on plant-herbivore interactions focuses on plant trait mean values, but recent studies showing the importance of plant genetic diversity for herbivores suggest that plant trait variance may be equally important. The consequences of plant trait variance for herbivore performance, however, have been largely overlooked. Here we report an extensive assessment of the effects of within-population plant trait variance on herbivore performance using 457 performance datasets from 53 species of insect herbivores. We show that variance in plant nutritive traits substantially reduces mean herbivore performance via non-linear averaging of performance relationships that were overwhelmingly concave down. By contrast, relationships between herbivore performance and plant defence levels were typically linear, with variance in plant defence not affecting herbivore performance via non-linear averaging. Our results demonstrate that plants contribute to the suppression of herbivore populations through variable nutrient levels, not just by having low average quality as is typically thought. We propose that this phenomenon could play a key role in the suppression of herbivore populations in natural systems, and that increased nutrient heterogeneity within agricultural crops could contribute to the sustainable control of insect pests in agroecosystems.
177 citations
Journal Article•
TL;DR: In this article, a review of studies of ungulate effects on plant community composition and ecosystem processes is presented, based on the premise that the effect ungulates exert on plant communities depends on the balance between feeding selectivity of herbivores and differences among plant species in their ability to recover from tissue loss.
Abstract: Large mammalian herbivores not only depend on plant communities for their existence but cause major changes in plant community composition and structure. These changes have direct consequences for ecosystem processes, but recent studies of ungulate-ecosystem relations show widely divergent ungulate effects in different ecosystems. We reviewed studies of ungulate effects on plant community composition to gain insight into potential mechanisms of ungulate-induced changes in both community composition and ecosystem processes. Our analysis of these studies is based on the premise that the effect ungulates exert on plant communities depends on the balance between (1) feeding selectivity of herbivores (i.e., degree to which dif- ferent plant species or ecotypes experience different levels of tissue loss), and (2) differences among plant species in their ability to recover from tissue loss. A large number of studies clearly show that selective ungulate herbivory leads to the dominance of unpalatable, chemically defended plant species in communities. However, many studies have also demonstrated that intensive long-term herbivory does not lead to the invasion of unpalatable species into the community, and can even increase the dominance of highly palatable species. Our review indicates that high levels of nutrient inputs or recycling and an intermittent temporal pattern of her- bivory (often due to migration) are key factors increasing the regrowth capacity of palatable species and hence maintaining their dominance in plant communities supporting abundant herbivores. Key factors limiting un- gulate foraging selectivity, again limiting herbivore-induced dominance of slow-growing, unpalatable species, include herding behavior, early growing season and postfire herbivory, asynchronous phenology of palatable versus unpalatable species, and low relative abundance of unpalatable species. Our review indicates differences among ecosystems in the role played by ungulate herbivory result from the relative strength of these factors enhancing plant tolerance to herbivory and limiting foraging selectivity. Anthropogenic changes in these factors (e.g., alteration of migration patterns) therefore have the potential to significantly alter the effects of ungulates on plant communities and ecosystem processes.
114 citations
TL;DR: It is proposed to exploit natural variation among plant species to assess the ecological costs of plant responses to herbivores and pollinators and to test if indirect plant resistance can conflict with pollinator attraction.
113 citations
TL;DR: Root defences must be investigated in their own ecophysiological and evolutionary context, specifically one which incorporates root microbial symbionts and antagonists, if the authors are to better understand the battle between plants and their hidden herbivores.
Abstract: The distinctive ecology of root herbivores, the complexity and diversity of root-microbe interactions, and the physical nature of the soil matrix mean that plant responses to root herbivory extrapolate poorly from our understanding of responses to aboveground herbivores. For example, root attack induces different changes in phytohormones to those in damaged leaves, including a lower but more potent burst of jasmonates in several plant species. Root secondary metabolite responses also differ markedly, although patterns between roots and shoots are harder to discern. Root defences must therefore be investigated in their own ecophysiological and evolutionary context, specifically one which incorporates root microbial symbionts and antagonists, if we are to better understand the battle between plants and their hidden herbivores.
111 citations
TL;DR: The experiments demonstrate that indole directly decreases food consumption, plant damage and survival of S. littoralis caterpillars, and shows that the effects of a plant secondary metabolite on weight gain and food consumption can diverge in a counterintuitive manner, which implies that larval growth can be a poor proxy for herbivore fitness and plant resistance.
Abstract: Plants release herbivore-induced volatiles (HIPVs), which can be used as cues by plants, herbivores and natural enemies. Theory predicts that HIPVs may initially have evolved because of their direct benefits for the emitter and were subsequently adopted as infochemicals.
Here, we investigated the potential direct benefits of indole, a major HIPV constituent of many plant species and a key defence priming signal in maize. We used indole-deficient maize mutants and synthetic indole at physiologically relevant doses to document the impact of the volatile on the generalist herbivore Spodoptera littoralis.
Our experiments demonstrate that indole directly decreases food consumption, plant damage and survival of S. littoralis caterpillars. Surprisingly, exposure to volatile indole increased caterpillar growth. Furthermore, we show that S. littoralis caterpillars and adults consistently avoid indole-producing plants in olfactometer experiments, feeding assays and oviposition trials.
Synthesis. Together, these results provide a potential evolutionary trajectory by which the release of a HIPV as a direct defence precedes its use as a cue by herbivores and an alert signal by plants. Furthermore, our experiments show that the effects of a plant secondary metabolite on weight gain and food consumption can diverge in a counterintuitive manner, which implies that larval growth can be a poor proxy for herbivore fitness and plant resistance.
82 citations
TL;DR: Coleoptera, the largest and the most diverse Insecta order, is characterized by multiple adaptations to plant feeding, and various aspects of the interactions among insects, microbes, and plants are described with a focus on coleopteran species, their bacterial symbionts, and their plant hosts to demonstrate that many factors contribute to the success of colepteran herbivory.
Abstract: Coleoptera, the largest and the most diverse Insecta order, is characterized by multiple adaptations to plant feeding Insect-associated microorganisms can be important mediators and modulators of interactions between insects and plants
Interactions between plants and insects are highly complex and involve multiple factors There are various defense mechanisms initiated by plants upon attack by herbivorous insects, including the development of morphological structures and the synthesis of toxic secondary metabolites and volatiles In turn, herbivores have adapted to feeding on plants and further sophisticated adaptations to overcome plant responses may continue to evolve Herbivorous insects may detoxify toxic phytocompounds, sequester poisonous plant factors, and alter their own overall gene expression pattern Moreover, insects are associated with microbes, which not only considerably affect insects, but can also modify plant defense responses to the benefit of their host Plants are also frequently associated with endophytes, which may act as bioinsecticides Therefore, it is very important to consider the factors influencing the interaction between plants and insects Herbivorous insects cause considerable damage to global crop production Coleoptera is the largest and the most diverse order in the class Insecta In this review, various aspects of the interactions among insects, microbes, and plants are described with a focus on coleopteran species, their bacterial symbionts, and their plant hosts to demonstrate that many factors contribute to the success of coleopteran herbivory
70 citations
TL;DR: Overall, plant tolerance and chemical defence to herbivores are affected by AMF through changes in basic plant physiology such as nutrient status, allocation patterns and growth rate, and this study suggests that it may be possible to predict effects of AMF on plant defences through these simple traits.
Abstract: Summary
Arbuscular mycorrhizal fungi (AMF) are ubiquitous plant symbionts that affect plant nutrient status, patterns of resource allocation and rates of plant growth. In addition, AMF influence the expression of plant defence, which may affect subsequent interactions between plants and higher trophic levels. Tolerance to herbivory and chemical defence represent two distinct defence strategies of plants, and there is accumulating evidence that AMF can influence each strategy separately. However, investigations on the simultaneous effects of AMF on tolerance and chemical defence mechanisms are lacking.
Using six milkweed (Asclepias) species and three levels of AMF inoculum, we show that plant tolerance to herbivory increases with foliar P concentrations, while both foliar cardenolide concentration and latex exudation (major chemical defences in milkweeds) increase with foliar N concentrations, but decrease with plant growth rate. Additionally, foliar cardenolide concentration increases with foliar P concentration, and latex exudation increases with root biomass. Because AMF have significant effects on all of these traits (foliar N and P concentrations, root biomass and growth rate), AMF can change plant tolerance and chemical defence simultaneously, albeit through different mechanisms.
Synthesis. Overall, plant tolerance and chemical defence to herbivores are affected by AMF through changes in basic plant physiology such as nutrient status, allocation patterns and growth rate, and our study suggests that it may be possible to predict effects of AMF on plant defences through these simple traits.
64 citations
TL;DR: Results indicate that the benefits of deterring insect herbivores by signalling may be balanced by the higher photosynthetic rate of non-signalling plants, a novel mechanism for the maintenance of leaf colour polymorphisms in nature.
Abstract: Summary
Whether plants use leaf colour to deter herbivores remains controversial. The warning signal hypothesis predicts that red pigmentation is adaptive by reducing herbivory; plants with predominantly red foliage should have higher fitness than those with green leaves. Despite many discussions, this prediction has rarely been tested, and alternative, non-exclusive hypotheses cannot be ruled out.
We have exploited leaf colour polymorphism in Pseudowintera colorata to test the warning signal hypothesis and to address possible alternative explanations.
Consistent with warning signals, redder foliage contained higher concentrations of polygodial, a sesquiterpene dialdehyde with strong antifeedant properties, and incurred less herbivory than green leaves. Redder plants hosted 22% fewer lepidopteran leafroller larvae than neighbouring green plants.
However, contrary to the predictions of the hypothesis, there were no differences in fitness parameters between red and green plants. Overall leaf canopy colour was not a significant predictor of the number of seeds per fruit or of mean seed weight. This may be explained by differences in photosynthesis: green P. colorata leaves had 47% higher maximum CO2 assimilation rates than matched red leaves from neighbouring plants.
These results indicate that the benefits of deterring insect herbivores by signalling may be balanced by the higher photosynthetic rate of non-signalling plants. A balance between signalling and photosynthesis is a novel mechanism for the maintenance of leaf colour polymorphisms in nature.
Synthesis. Anthocyanin pigments may simultaneously serve multiple functions within leaves, and individuals of the same plant species may use different strategies to cope with insect herbivores. Therefore, investigations into the role of these pigments in plant–insect interactions need to consider plant physiology and the diversity of plant defence mechanisms.
60 citations
TL;DR: It is suggested that native generalist herbivores promote enemy release of some plant invaders by avoiding them and contribute to biotic resistance of others by consuming them, and herbivore selectivity likely plays an important role in the invasion process.
Abstract: Native generalist herbivores might limit plant invasion by consuming invading plants or enhance plant invasion by selectively avoiding them. The role of herbivores in plant invasion has been investigated in relation to plant native/introduced status, however, a knowledge gap exists about whether food selection occurs according to native/introduced status or to species. We tested preference of the native herbivore white-tailed deer (Odocoileus virginianus) for widespread and frequently occurring invasive introduced and native plants in the northeastern United States. Multiple-choice deer preference trials were conducted for the species and relative preference was determined using biomass consumption and feeding behavior. While more native than introduced plant biomass was consumed overall, deer food selection varied strongly by plant species. Results show consistent deer avoidance of several invasive introduced plants (Alliaria petiolata, Berberis thunbergii, and Microstegium vimineum) and a native plant (Dennstaedtia punctilobula). Other invasive introduced plants (Celastrus orbiculatus, Ligustrum vulgare, and Lonicera morrowii) and a native plant (Acer rubrum) were highly preferred. These results provide evidence that herbivore impacts on plant invaders depend on plant species palatability. Consequently, herbivore selectivity likely plays an important role in the invasion process. To the extent that herbivory impacts population demographics, these results suggest that native generalist herbivores promote enemy release of some plant invaders by avoiding them and contribute to biotic resistance of others by consuming them.
53 citations
TL;DR: Fitness costs of glyphosate-resistant plants were expressed under both conditions of water availability and could be useful for designing management strategies and exploiting the reduced glyphosate- resistant perennial ryegrass fitness in the absence of glyphosate selection.
Abstract: The net selection effect of herbicides on herbicide-resistance traits in weeds is conditioned by the fitness benefits and costs associated with resistance alleles. Fitness costs play an important evolutionary role preventing the fixation of adaptive alleles and contributing to the maintenance of genetic polymorphisms within populations. Glyphosate is widely used in world agriculture, which has led to the evolution of widespread glyphosate resistance in many weed species. The fitness of glyphosate-resistant and -susceptible perennial ryegrass plants selected from within a single population were studied in two field experiments conducted during 2011 and 2012 under different soil water availability. Glyphosate-resistant plants showed a reduction in height of 12 and 16%, leaf blade area of 16 and 33%, shoot biomass of 45 and 55%, seed number of 33 and 53%, and total seed mass of 16 and 5% compared to glyphosate-susceptible plants in 2011 and 2012, respectively. The reduction in seed number per plant ...
TL;DR: It is demonstrated that jasmonate-deficient Nicotiana attenuata plants suffer more damage by arthropod and vertebrate herbivores than jasMonate-producing plants in nature, and Ecological and evolutionary models on plant defense signaling should aim at integrating arthropOD and vertebrates herbivory at the community level.
Abstract: Endogenous jasmonates are important regulators of plant defenses. If and how they enable plants to maintain their reproductive output when facing community-level herbivory under natural conditions, however, remains unknown. We demonstrate that jasmonate-deficient Nicotiana attenuata plants suffer more damage by arthropod and vertebrate herbivores than jasmonate-producing plants in nature. However, only damage by vertebrate herbivores translates into a significant reduction in flower production. Vertebrate stem peeling has the strongest negative impact on plant flower production. Stems are defended by jasmonate-dependent nicotine, and the native cottontail rabbit Sylvilagus nuttallii avoids jasmonate-producing N. attenuata shoots because of their high levels of nicotine. Thus, endogenous jasmonates enable plants to resist different types of herbivores in nature, and jasmonate-dependent defenses are important for plants to maintain their reproductive potential when facing vertebrate herbivory. Ecological and evolutionary models on plant defense signaling should aim at integrating arthropod and vertebrate herbivory at the community level.
TL;DR: This study is the first to demonstrate that parasitoids, through increasing feeding by their herbivorous hosts, can induce higher levels of non-volatile plant chemical defenses.
Abstract: Insect parasitoids can play ecologically important roles in virtually all terrestrial plant–insect herbivore interactions, yet whether parasitoids alter the defensive traits that underlie interactions between plants and their herbivores remains a largely unexplored question. Here, we examined the reciprocal trophic interactions among populations of the wild cabbage Brassica oleracea that vary greatly in their production of defensive secondary compounds – glucosinolates (GSs), a generalist herbivore, Trichoplusia ni, and its polyembryonic parasitoid Copidosoma floridanum. In a greenhouse environment, plants were exposed to either healthy (unparasitized), parasitized, or no herbivores. Feeding damage by herbivores induced higher levels of the indole GSs, glucobrassicin and neoglucobrassicin, but not any of the other measured GSs. Herbivores parasitized by C. floridanum induced cabbage plants to produce 1.5 times more indole GSs than levels induced by healthy T. ni and five times more than uninduced plants. As a gregarious endoparasitoid, C. floridanum causes its host T. ni to feed more than unparasitized herbivores resulting in increased induction of indole GSs. In turn, herbivore fitness parameters (including differential effects on male and female contributions to lifetime fecundity in the herbivore) were negatively correlated with the aliphatic GSs, sinigrin and gluconapin, whereas parasitoid fitness parameters were negatively correlated with the indole GSs, glucobrassicin and neoglucobrassicin. That herbivores and their parasitoids appear to be affected by different sets of GSs was unexpected given the intimate developmental associations between host and parasitoid. This study is the first to demonstrate that parasitoids, through increasing feeding by their herbivorous hosts, can induce higher levels of non-volatile plant chemical defenses. While parasitoids are widely recognized to be ubiquitous in most terrestrial insect herbivore communities, their role in influencing plant–insect herbivore relationships is still vastly underappreciated.
TL;DR: Observed evolutionary convergence for multiple plant traits, between the natural and manipulative experiments, emphasizes the role of insect herbivores as key drivers of plant adaptation and geographic differentiation.
Abstract: Summary
Herbivory can drive rapid evolution of plant chemical traits mediating defensive and competitive ability. At a geographic scale, plant populations that escape selection from their ancestral herbivores may evolve decreased defence and increased competitiveness. While contrasts between native and invasive populations of plants lend support to this hypothesis, such experiments cannot establish causal links between herbivory and evolved invasive phenotypes.
Here, we conducted geographic contrasts, and coupled these with long-term selection experiments that directly test for evolutionary responses to herbivore exclusion. In common gardens, we contrasted Solidago altissima genotypes that were historically exposed or protected from herbivory across two experimental time-scales: (i) a natural experiment where plant populations evolved either with native herbivory (in Minnesota and New York) or evolved relatively free from herbivory for ˜100 years in Japan, and (ii) a 12-year manipulative experiment where plants were either exposed to ambient herbivory or treated with insecticide.
In both experiments, plant populations responded to herbivore release by evolving increased production of root allelochemicals and interspecific competitive ability against Poa pratensis. While plant resistance to a beetle herbivore did not diverge between plant origins, we still observed parallel evolutionary shifts in leaf secondary metabolite and protease inhibitor production, which may confer resistance to diverse herbivore species.
Synthesis. Observed evolutionary convergence for multiple plant traits, between the natural and manipulative experiments, emphasizes the role of insect herbivores as key drivers of plant adaptation and geographic differentiation.
TL;DR: Assessment of the effects of single or dual herbivory by Brevicoryne brassicae aphids and/or Plutella xylostella caterpillar on resistance of plants from three natural populations of wild cabbage to feeding by caterpillars of Mamestra brassicAE contributes to the understanding of plant defence plasticity in response to multiple insect attacks.
Abstract: Plants are part of biodiverse communities and frequently suffer from attack by multiple herbivorous insects. Plant responses to these herbivores are specific for insect feeding guilds: aphids and caterpillars induce different plant phenotypes. Moreover, plants respond differentially to single or dual herbivory, which may cascade into a chain of interactions in terms of resistance to other community members. Whether differential responses to single or dual herbivory have consequences for plant resistance to yet a third herbivore is unknown. We assessed the effects of single or dual herbivory by Brevicoryne brassicae aphids and/or Plutella xylostella caterpillars on resistance of plants from three natural populations of wild cabbage to feeding by caterpillars of Mamestra brassicae. We measured plant gene expression and phytohormone concentrations to illustrate mechanisms involved in induced responses. Performance of both B. brassicae and P. xylostella was reduced when feeding simultaneously with the other herbivore, compared to feeding alone. Gene expression and phytohormone concentrations in plants exposed to dual herbivory were different from those found in plants exposed to herbivory by either insect alone. Plants previously induced by both P. xylostella and B. brassicae negatively affected growth of the subsequently arriving M. brassicae. Furthermore, induced responses varied between wild cabbage populations. Feeding by multiple herbivores differentially activates plant defences, which has plant-mediated negative consequences for a subsequently arriving herbivore. Plant population-specific responses suggest that plant populations adapt to the specific communities of insect herbivores. Our study contributes to the understanding of plant defence plasticity in response to multiple insect attacks.
TL;DR: In the field, early-season herbivory negatively affected plant reproduction and seeds of these plants suffered lower levels of infestation by seed-eating beetles, which in turn suffered less parasitism, which provided insight into how direct and indirect interactions between and within different trophic levels affect the dynamics and structure of complex communities.
Abstract: There is an increasing awareness that herbivory by one insect species induces changes in a plant that affect the performance of other herbivore species that feed on the same plant. However, previous studies of interspecies interactions mediated by plant defense responses have rarely taken into account different insect guilds or the third trophic level. Using a combination of field and laboratory experiments, we examined how early-season herbivory in lima bean plants (Phaseolus lunatus) by the leaf-chewing herbivore Cerotoma ruficornis and the bean pod weevil Apion godmani affects the abundance and performance of the seed beetle Zabrotes subfasciatus and that of its parasitoid Stenocorse bruchivora, which occurs on the plants at the end of the growing season. In addition, we determined the consequences of early-season herbivore-induced defenses on plant performance. We hypothesized that early-season induction would affect plant reproduction and, hence, would alter the suitability of seeds for late-season seed-eating beetles, and that this would in turn alter the vulnerability of these seed beetles to parasitoids. We found strong support for these hypotheses. In the field, early-season herbivory negatively affected plant reproduction and seeds of these plants suffered lower levels of infestation by seed-eating beetles, which in turn suffered less parasitism. Laboratory assays with field-collected seeds confirmed that the performance of beetles and parasitoids was lower on seeds from plants that had been subjected to early-season herbivory. Further analyses revealed that seeds produced by control plants were larger, heavier, and had a higher concentration of cyanogenic glycosides and total protein content than seeds from plants subjected to herbivory. Our results provide insight into how direct and indirect interactions between and within different trophic levels affect the dynamics and structure of complex communities.
TL;DR: Larval feeding of an aquatic insect, the moth Parapoynx stratiotata, showed strong feeding preferences demonstrating that aquatic insects can directly and indirectly alter water quality and vegetation composition.
Abstract: Eutrophication and globalisation facilitate the dominance of exotic plants in aquatic ecosystems worldwide. Aquatic omnivores can provide biotic resistance to plant invasions, but little is known about whether obligate aquatic herbivores can do the same. Herbivores such as insects can decimate aquatic vegetation, but may not be able to consume exotic plants due to their more or less specialised nature of feeding. We experimentally tested the larval feeding of an aquatic insect, the moth Parapoynx stratiotata, on eleven submerged plant species, from either native or exotic origin. We also tested whether insect herbivory stimulates nutrient and organic matter release, thus affecting water quality. Larvae of P. stratiotata consumed seven out of eleven plant species, and their growth was related to plant nutrient content and stoichiometry. However, larvae had no preference for either native or exotic macrophytes, and their plant preference was not related to the measured plant traits, but was possibly driven by secondary metabolites. Through plant consumption, caterpillars induced brownification and phosphate release, and the intensity thereof varied among plant species, but not between native and exotic plants. In conclusion, P. stratiotata showed strong feeding preferences demonstrating that aquatic insects can directly and indirectly alter water quality and vegetation composition.
TL;DR: The plant variability-gut acclimation hypothesis is proposed, which posits that plant chemical variability constrains herbivore anti-predator defenses by frequently requiring herbivores to acclimate their guts to changing plant defenses and nutrients.
Abstract: Variability in plant chemistry has long been believed to suppress populations of insect herbivores by constraining herbivore resource selection behavior in ways that make herbivores more vulnerable to predation. The focus on behavior, however, overlooks the pervasive physiological effects of plant variability on herbivores. Here we propose the plant variability-gut acclimation hypothesis, which posits that plant chemical variability constrains herbivore anti-predator defenses by frequently requiring herbivores to acclimate their guts to changing plant defenses and nutrients. Gut acclimation, including changes to morphology and detoxification enzymes, requires time and nutrients, and we argue these costs will constrain how and when herbivores can mount anti-predator defenses. A consequence of this hypothesis is stronger top-down control of herbivores in heterogeneous plant populations.
TL;DR: Avoidance of mildew-infested plants may be adaptive for C. glomerata parasitoids, whereas P. brassicae caterpillars may suffer less parasitism on mildEW-infected plants in nature.
Abstract: The co-occurrence of different antagonists on a plant can greatly affect infochemicals with ecological consequences for higher trophic levels. Here we investigated how the presence of a plant pathogen, the powdery mildew Erysiphe cruciferarum, on Brassica rapa affects (1) plant volatiles emitted in response to damage by a specialist herbivore, Pieris brassicae; (2) the attraction of the parasitic wasp Cotesia glomerata and (3) the performance of P. brassicae and C. glomerata. Plant volatiles were significantly induced by herbivory in both healthy and mildew-infected plants, but were quantitatively 41% lower for mildew-infected plants compared to healthy plants. Parasitoids strongly preferred Pieris-infested plants to dually-infested (Pieris + mildew) plants, and preferred dually infested plants over only mildew-infected plants. The performance of P. brassicae was unaffected by powdery mildew, but C. glomerata cocoon mass was reduced when parasitized caterpillars developed on mildew-infected plants. Thus, avoidance of mildew-infested plants may be adaptive for C. glomerata parasitoids, whereas P. brassicae caterpillars may suffer less parasitism on mildew-infected plants in nature. From a pest management standpoint, the concurrent presence of multiple plant antagonists can affect the efficiency of specific natural enemies, which may in turn have a negative impact on the regulation of pest populations.
TL;DR: It is demonstrated that salt stress in Indian mustard lessens the effect of herbivory on plant mass, and increases the plants' constitutive resistance to herbivores.
Abstract: Plants often grow under the combined stress of several factors Salinity and herbivory, separately, can severely hinder plant growth and reproduction, but the combined effects of both factors are still not clearly understood Salinity is known to reduce plant tissue nitrogen content and growth rates Since herbivores prefer tissues with high N content, and biochemical pathways leading to resistance are commonly elicited by salt-stress, we hypothesized that plants growing in saline conditions would have enhanced resistance against herbivores The non-halophyte, Brassica juncea, and the generalist herbivore Trichoplusia ni were used to test the prediction that plants subjected to salinity stress would be both more resistant and more tolerant to herbivory than those growing without salt stress Plants were grown under different NaCl levels, and either exposed to herbivores and followed by removal of half of their leaves, or left intact Plants were left to grow and reproduce until senescence Tissue quality was assessed, seeds were counted and biomass of different organs measured Plants exposed to salinity grew less, had reduced tissue nitrogen, protein and chlorophyll content, although proline levels increased Specific leaf area, leaf water content, transpiration and root:shoot ratio remained unaffected Plants growing under saline condition had greater constitutive resistance than unstressed plants However, induced resistance and tolerance were not affected by salinity These results support the hypothesis that plants growing under salt-stress are better defended against herbivores, although in B juncea this may be mostly through resistance, and less through tolerance
TL;DR: This study suggests that an ecosystem engineer could have significant impact on herbivore community not only by changing plant morphology, but also by altering host quality and modifying plant-induced responses to subsequent herbivory.
Abstract: * Herbivores that modify plant morphology, such as gall-forming insects, can disproportionately impact arthropod community on their host plants by providing novel habitats and shelters from biotic and abiotic stresses. These ecosystem engineers could also modify plant chemical properties, but how such changes in plant quality affect the behaviour of subsequent colonizers has rarely been investigated. * We explored how an initial infestation of the tall goldenrod (Solidago altissima) by an ecosystem engineer, the rosette gall-midge (Rhopalomyia solidaginis), affects colonization behaviour of a shelter-using beetle (Microrhopala vittata) through plant-induced responses in the field. * Beetles preferentially colonized plants with galls and exhibited a clumped distribution on those plants, which suggested a possible advantage for aggregating on galled plants. Accordingly, we found that beetles remained longer on galled plants with previous beetle damage than those without beetle damage. No such effect of beetle damage was found on plants without a gall. Similar interactions between galler infestation and beetle damage were found in beetle's feeding choice, leaf diterpene and serine protease inhibitor production, and volatile organic compound (VOC) emission. These plant metabolic induction and herbivore response patterns indicated that the gall-midge can alter how plants respond to the beetle damage and that gall presence coupled with beetle damage improves leaf palatability for the beetle. Finally, we found reciprocal effects of beetles on gall-midge performance to be neutral to slightly positive, suggesting that the observed field association of the two herbivores could be formed by plant-mediated facilitation. * Synthesis. Our study suggests that an ecosystem engineer could have significant impact on herbivore community not only by changing plant morphology, but also by altering host quality and modifying plant-induced responses to subsequent herbivory. As such, R. solidaginis also functions as a keystone herbivore that has disproportionate effects on community dynamics and composition meditated by induced plant growth and metabolic responses.
TL;DR: It is found that overcompensation can cause bounded fluctuations in the herbivore density by itself, whereas neither tolerance nor induced resistance can cause an outbreak on its own.
Abstract: Outbreaks of herbivorous insects are detrimental to natural and agricultural systems, but the mechanisms driving outbreaks are not well understood. Plant responses to herbivory have the potential to produce outbreaks, but long-term effects of plant responses on herbivore dynamics are understudied. To quantify these effects, we analyze mathematical models of univoltine herbivores consuming annual plants with two responses: (1) compensatory regrowth, which affects herbivore survival in food-limited situations by increasing the amount of food available to the herbivore; and (2) induced resistance, which reduces herbivore survival proportional to the strength of the response. Compensatory regrowth includes tolerance, where plants replace some or all of the consumed biomass, and overcompensation, where plants produce more biomass than was consumed. We found that overcompensation can cause bounded fluctuations in the herbivore density (called outbreaks here) by itself, whereas neither tolerance nor indu...
TL;DR: It is found that aphid induced defenses and apparent competition combined such that plant damage by caterpillars was lowest when predators were present at low aphid density, which suggests that herbivores can benefit plants, but the effect on host plants is mediated by herbivore density.
Abstract: Herbivore-induced plant resistance and apparent competition are two indirect ways herbivores interact. If a less damaging herbivore indirectly suppresses the abundance of a more damaging herbivore via these mechanisms, then plants may ultimately benefit. Changes in herbivore density, however, can dictate the intensity of species interactions and may play a critical role in determining the outcome of plant- and predator-mediated herbivore interactions. We tested the effects of herbivore density on the strength of indirect interactions among phloem-feeding aphids and herbivorous caterpillars and the outcome of these interactions for their shared host plant, cotton (Gossypium hirsutum). We quantified the survival of caterpillars on host plants that were infested with varying densities of aphids in the presence and absence of predators (ladybeetles). We found that aphids induced defensive proteins in cotton plants and that caterpillar survival was negatively affected by induced resistance. Likewise, we found that the presence of aphids increased predation of caterpillars by ladybeetles, but that apparent competition between aphids and caterpillars was density dependent. Ladybeetles consumed relatively high numbers of small caterpillars at low to intermediate aphid densities, but essentially became aphid specialists at high aphid densities. Aphid induced defenses and apparent competition combined such that plant damage by caterpillars was lowest when predators were present at low aphid density (induced resistance + highest level of apparent competition). This suggests that herbivores can benefit plants, but the effect on host plants is mediated by herbivore density. Indirect herbivore-plant mutualisms may increase plant quality, plant fitness, and yield of crop plants and these interactions need to be considered in ecologically based pest management plans. In addition, these interactions likely alter arthropod community structure and natural selection on anti-herbivore defense traits in plants in natural systems.
TL;DR: Host-specific herbivores of different guilds showed similar responses to drought-stressed, fast-growing plants, which could discourage the attack of specialist insects in both sap-sucking and leaf-chewing feeding guilds.
Abstract: An initial destabilization of functions triggered by drought stress in plants is followed by acclimatization and acquisition of tolerance; however, knowledge remains limited on drought-mediated changes in plant quality for herbivores. We tested whether a water-stressed fast-growing plant negatively affects host-specialist insects in both sap-sucking and leaf-chewing feeding guilds. Collards (Brassica oleracea var. acephala) were grown in well-watered, slightly water-stressed and severely water-stressed conditions. Decreasing soil moisture adversely affected plant development, assessed as a reduction in leaf number and size, stomatal size and relative water content. Severely stressed plants had less fiber and glucosinolates; however, they showed more total nitrogen and lipids. Larval survival, pupal weight, reproductive rate (Ro) and rate of population growth (r) were lower when the leaf-chewing Plutella xylostella was reared with severely stressed collards. In multiple-choice tests, moths laid fewer eggs on leaf discs of collard that were exposed to drought. The fecundity of the sap-sucking Brevicoryne brassicae was higher and the development of alates was lower when insects were fed on plants kept in well-watered regime as compared to slight-stress and severe-stress. Despite higher nitrogen content and fewer glucosinolates, a higher level of leaf surface wax in severely stressed collards possibly decreased food quality for both herbivores. Thus, host-specific herbivores of different guilds showed similar responses to drought-stressed, fast-growing plants. Water-stressed crops could discourage the attack of specialist insects, but the intensity of the stress that is required to achieve this effect will greatly reduce crop production, in terms of plant growth or foliage increment.
TL;DR: It is concluded that plant cuttings can serve as adequate proxies for live plants for preference/performance studies, but that experimentalists should be aware of the variation imposed both by plant handling and plant phenology for female oviposition preference.
Abstract: Plants have evolved a number of defences to ameliorate herbivore attacks including chemicals induced by mechanical wounding. Such changes in plant chemical composition are potential confounding factors in experiments on plant–insect interactions, which often present cuttings of potential host plants to phytophagous insects. In particular, this could affect studies of female egg-laying preference and larval performance, because the same plant chemicals that deter certain generalist insects can elevate attacks from more specialized insects. Furthermore, plant cuttings are by definition smaller than intact plants, and any female host size preference could thus affect experiments using plant cuttings. We first assessed female preference and larval performance of a specialist herbivore, Pieris napi (L.) (Lepidoptera: Pieridae, Pierini), confronted with either intact plants or leaf-cuttings of four Brassicaceae host plants, Alliaria petiolata (Bieb.) Cavara & Grande, Barbarea vulgaris (L.) WT Aiton, Berteroa incana (L.) DC., and Brassica napus (L.). Egg and larval survival did not differ between intact plants and leaf-cuttings, whereas larval growth was slightly, but significantly, faster on leaf-cuttings. Females, however, significantly preferred to lay eggs on intact plants of all four hosts, although the preference hierarchy for the intact plants was largely mirrored by that for leaf-cuttings. We then tested the female preference for different size-classes of intact B. napus plants. Small individuals received more eggs than larger individuals, and follow-up experiments showed that this difference was largely generated by a strong female preference for cotyledon leaves; there was no significant difference in female preference for large and small individuals when both carried cotyledons, and females landing on cotyledons were more likely to oviposit compared to when landing on a true leaf. Our study concludes that plant cuttings can serve as adequate proxies for live plants for preference/performance studies, but that experimentalists should be aware of the variation imposed both by plant handling and plant phenology for female oviposition preference.
TL;DR: Plant defence can be understood fully when herbivores and their victims are considered in the broader context of other interactions among coexisting species and of the medium in which these interactions occur.
TL;DR: The results show plant-plant signalling: ‘Nyamula’ maize plants emitting oviposition-induced volatiles attractive to the herbivore’s natural enemies can induce this indirect defence trait in conspecific neighbouring undamaged maize plants.
Abstract: Attack of plants by herbivorous arthropods may result in considerable changes to the plant’s chemical phenotype with respect to emission of herbivore-induced plant volatiles (HIPVs). These HIPVs have been shown to act as repellents to the attacking insects as well as attractants for the insects antagonistic to these herbivores. Plants can also respond to HIPV signals from other plants that warn them of impending attack. Recent investigations have shown that certain maize varieties are able to emit volatiles following stemborer egg deposition. These volatiles attract the herbivore’s parasitoids and directly deter further oviposition. However, it was not known whether these oviposition-induced maize (Zea mays, L.) volatiles can mediate chemical phenotypic changes in neighbouring unattacked maize plants. Therefore, this study sought to investigate the effect of oviposition-induced maize volatiles on intact neighbouring maize plants in ‘Nyamula’, a landrace known to respond to oviposition, and a standard commercial hybrid, HB515, that did not. Headspace volatile samples were collected from maize plants exposed to Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) egg deposition and unoviposited neighbouring plants as well as from control plants kept away from the volatile emitting ones. Behavioural bioassays were carried out in a four-arm olfactometer using egg (Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)) and larval (Cotesia sesamiae Cameron (Hymenoptera: Braconidae)) parasitoids. Coupled Gas Chromatography-Mass Spectrometry (GC-MS) was used for volatile analysis. For the ‘Nyamula’ landrace, GC-MS analysis revealed HIPV production not only in the oviposited plants but also in neighbouring plants not exposed to insect eggs. Higher amounts of EAG-active biogenic volatiles such as (E)-4,8-dimethyl-1,3,7-nonatriene were emitted from these plants compared to control plants. Subsequent behavioural assays with female T. bournieri and C. sesamiae parasitic wasps indicated that these parasitoids preferred volatiles from oviposited and neighbouring landrace plants compared to those from the control plants. This effect was absent in the standard commercial hybrid we tested. There was no HIPV induction and no difference in parasitoid attraction in neighbouring and control hybrid maize plants. These results show plant-plant signalling: ‘Nyamula’ maize plants emitting oviposition-induced volatiles attractive to the herbivore’s natural enemies can induce this indirect defence trait in conspecific neighbouring undamaged maize plants. Maize plants growing in a field may thus benefit from this indirect defence through airborne signalling which may enhance the fitness of the volatile-emitting plant by increasing predation pressure on herbivores.
TL;DR: Plants damaged by herbivores emit blends of volatile organic compounds (VOCs) that attract the herbivore’s natural enemies, and whether attraction of the natural enemies that responded to aphid-induced VOCs was altered by simultaneous damage by a chewing Herbivore is investigated.
Abstract: Plants damaged by herbivores emit blends of volatile organic compounds (VOCs) that attract the herbivore’s natural enemies. Most work has focussed on systems involving one plant, one herbivore and one natural enemy, though, in nature, plants support multiple herbivores and multiple natural enemies of these herbivores. Our study aimed to understand how different aphid natural enemies respond to aphid-induced VOCs, and whether attraction of the natural enemies that responded to aphid-induced VOCs was altered by simultaneous damage by a chewing herbivore. We used a model system based on Brassica juncea (Brassicaceae), Myzus persicae (Hemiptera: Aphididae) and Plutella xylostella (Lepidoptera: Plutellidae). Ceraeochrysa cubana (Neuroptera: Chrysopidae) did not show preferences for any plant odour, while Cycloneda sanguinea (Coleoptera: Coccinellidae) responded to undamaged plants over air but not to aphid-damaged plants over undamaged plants. Therefore, no further tests were carried out with these two species. Chrysoperla externa (Neuroptera: Chrysopidae) preferred aphid-damaged plants, but not caterpillar-damaged plants, over undamaged plants, and preferred plants damaged by both herbivores over both undamaged plants and aphid-damaged plants. When tested for responses against undamaged plants, Aphidius colemani (Hymenoptera: Braconidae) preferred aphid-damaged plants but not plants damaged by caterpillars. Plants damaged by both herbivores attracted more parasitoids than undamaged plants, but not more than aphid-damaged plants. Thus, multiply damaged plants were equally attractive to A. colemani and more attractive to C. externa than aphid-damaged plants, while C. cubana and C. sanguinea did not respond to aphid-induced VOCs, highlighting how different natural enemies can have different responses to herbivore-damaged plants.
TL;DR: Evidence that natal plants affect competitive interactions on another plant species is provided, and the potential importance of neighboring plant species on herbivore community composition in agricultral systems is highlighted.
Abstract: Interspecific competition between herbivores is widely recognized as an important determinant of community structure. Although researchers have identified a number of factors capable of altering competitive interactions, few studies have addressed the influence of neighboring plant species. If adaptation to/ epigenetic effects of an herbivore’s natal host plant alter its performance on other host plants, then interspecific herbivore interactions may play out differently in heterogeneous and homogenous plant communities. We tested wether the natal host plant of a whitefly population affected interactions between the Middle-east Asia Minor 1 (MEAM1) and Mediterranean (MED) cryptic species of the whitefly Bemisia tabaci by rearing the offspring of a cabbage-derived MEAM1 population and a poinsettia-derived MED population together on three different host plants: cotton, poinsettia, and cabbage. We found that MED dominated on poinsettia and that MEAM1 dominated on cabbage, results consistent with previous research. MED also dominated when reared with MEAM1 on cotton, however, a result at odds with multiple otherwise-similar studies that reared both species on the same natal plant. Our work provides evidence that natal plants affect competitive interactions on another plant species, and highlights the potential importance of neighboring plant species on herbivore community composition in agricultral systems.
TL;DR: This study shows that within a given plant species, plant‐mediated interactions between herbivores of the same feeding guild can be stable, which opens up the possibility of adaptations by associated organisms and suggests that plant‐ mediated interactions may contribute more strongly to evolutionary dynamics in terrestrial (agro)ecosystems than previously assumed.
Abstract: Plant-mediated interactions between herbivores are important determinants of community structure and plant performance in natural and agricultural systems. Current research suggests that the outcome of the interactions is determined by herbivore and plant identity, which may result in stochastic patterns that impede adaptive evolution and agricultural exploitation. However, few studies have systemically investigated specificity versus general patterns in a given plant system by varying the identity of all involved players. We investigated the influence of herbivore identity and plant genotype on the interaction between leaf-chewing and root-feeding herbivores in maize using a partial factorial design. We assessed the influence of leaf induction by oral secretions of six different chewing herbivores on the response of nine different maize genotypes and three different root feeders. Contrary to our expectations, we found a highly conserved pattern across all three dimensions of specificity: The majority of leaf herbivores elicited a negative behavioral response from the different root feeders in the large majority of tested plant genotypes. No facilitation was observed in any of the treatment combinations. However, the oral secretions of one leaf feeder and the responses of two maize genotypes did not elicit a response from a root-feeding herbivore. Together, these results suggest that plant-mediated interactions in the investigated system follow a general pattern, but that a degree of specificity is nevertheless present. Our study shows that within a given plant species, plant-mediated interactions between herbivores of the same feeding guild can be stable. This stability opens up the possibility of adaptations by associated organisms and suggests that plant-mediated interactions may contribute more strongly to evolutionary dynamics in terrestrial (agro)ecosystems than previously assumed.