Chewing Herbivores Research Articles

Herbivore-induced plant volatiles accurately predict history of coexistence, diet breadth, and feeding mode of herbivores.

Herbivore-induced plant volatiles (HIPVs) serve as specific cues to higher trophic levels. Novel, exotic herbivores entering native foodwebs may disrupt the infochemical network as a result of changes in HIPV profiles. Here, we analysed HIPV blends of native Brassica rapa plants infested with one of 10 herbivore species with different coexistence histories, diet breadths and feeding modes. Partial least squares (PLS) models were fitted to assess whether HIPV blends emitted by Dutch B.rapa differ between native and exotic herbivores, between specialists and generalists, and between piercing-sucking and chewing herbivores. These models were used to predict the status of two additional herbivores. We found that HIPV blends predicted the evolutionary history, diet breadth and feeding mode of the herbivore with an accuracy of 80% or higher. Based on the HIPVs, the PLS models reliably predicted that Trichoplusia ni and Spodoptera exigua are perceived as exotic, leaf-chewing generalists by Dutch B.rapa plants. These results indicate that there are consistent and predictable differences in HIPV blends depending on global herbivore characteristics, including coexistence history. Consequently, native organisms may be able to rapidly adapt to potentially disruptive effects of exotic herbivores on the infochemical network.

Not all droughts are created equal? The effects of stress severity on insect herbivore abundance

Predicting interactions between drought and plant–insect interactions has been a challenge. Currently, we are unable to accurately predict herbivore abundance on stressed plants despite over 500 publications and half a dozen formal hypotheses. With drought predicted to increase in severity with climate change, determining herbivore abundance on stressed plants is critical for continued agricultural and natural system management. During drought, plants increase concentrations of nutrients and also suffer from water loss. Many empirical studies test drought on plants using severe continuous stress, but studies suggest drought may benefit herbivores when it is intermittent (pulsed). In our study, we tested intermittent and severe stress on herbivore and arthropod abundance in a cotton agro-ecosystem. Our goal was to determine how these types of drought influence herbivore abundance on stressed plants and the relationship between herbivore abundance and stress-related nutrients. We found that intermittent and severe water-deficit stress had different effects on insect herbivores, signifying that drought severity influences herbivore abundance. Piercing–sucking herbivores such as thrips, stink bugs, and leafhoppers were more abundant on intermittently stressed plants than on severely stressed plants. Drought did not significantly affect chewing herbivores, and their abundance was inconsistent on stressed and well-watered plants. Furthermore, nutrient concentrations were similar between stressed and unstressed plants, but herbivore abundance was greater on intermittently stressed plants, suggesting that other physiological characteristics of stressed plants and herbivore feeding ecology must be considered. Our study suggests that the drought severity must be considered when predicting herbivore abundance on stressed plants.

Heavy metal (hyper)accumulation in leaves of Arabidopsis halleri is accompanied by a reduced performance of herbivores and shifts in leaf glucosinolate and element concentrations

Abiotic and biotic factors can significantly influence and modify the chemical composition of a plant. In this study the influences of exposure to cadmium and zinc and of herbivory on plant-insect interactions were investigated. We studied the performance of two herbivore species with different feeding modes (chewing versus sucking) on the metal-hyperaccumulator species Arabidopsis halleri (Brassicaceae) as well as individual glucosinolate (GLS) and element concentrations of young and old leaves in response to different soil and herbivore treatments. The performance of both insect species was reduced on plants grown on metal-amended soil compared to plants grown on unamended soil. This result supports the elemental defense hypothesis, which postulates that high internal metal concentrations can protect plants from herbivores and which was mostly tested with chewing herbivores so far. Alongside higher levels of heavy metals in plants grown on metal-amended soil, GLS and element concentrations were differentially affected in young versus old leaves and responded differently to herbivory. Concentrations of most GLS and elements were higher in young compared to old leaves, which is in line with the optimal defense theory, which predicts a better defense in more valuable tissue. Our results highlight that various organic as well as inorganic compounds in plants can be affected in different directions by the complex interplay between plants, herbivores, and heavy metal contamination of soil.

Herbivore‐specific induction of defence metabolites in a grass–endophyte association

Summary Plants have developed a variety of defence strategies against herbivores. One possible strategy is the induced production of metabolites following herbivore attack. Plant‐associated micro‐organisms can be the source of such defensive compounds. For example, cool‐season grasses can be associated with systemic endophytic fungi of the genus Epichloё, which produce herbivore‐toxic alkaloids. In a controlled common garden approach, we tested the hypothesis that different types of herbivory induce endophyte growth and increase the endophyte‐mediated production of three bioactive alkaloids which can deter or toxify herbivores. During 18 weeks, we analysed biweekly endophyte and alkaloid concentrations in the grass Lolium perenne infected with the endophytic fungus Epichloё festucae var. lolii. The experiment was conducted throughout the field season and compared three different herbivore treatments to the control treatment (herbivory exclosure). We showed that the concentration of the vertebrate toxic alkaloid lolitrem B increased following clipping (a simulation of grazing herbivores), while the insect deterring alkaloid peramine increased following locust herbivory (biting–chewing herbivores). The endophyte concentration increased slightly following clipping (P = 0·09). Sap sucking aphids altered neither endophyte nor alkaloid concentrations. Our study provides evidence for an herbivore‐specific induction of endophyte‐mediated responses following herbivore attack on its host grass. Our results suggest that the grass–endophyte symbiosis involves a close chemical crosstalk between the interacting partners.

Modulation of plant defense responses to herbivores by simultaneous recognition of different herbivore-associated elicitors in rice.

Induced plant defense responses against insect herbivores are triggered by wounding and/or perception of herbivore elicitors from their oral secretions (OS) and/or saliva. In this study, we analyzed OS isolated from two rice chewing herbivores, Mythimna loreyi and Parnara guttata. Both types of crude OS had substantial elicitor activity in rice cell system that allowed rapid detection of early and late defense responses, i.e. accumulation of reactive oxygen species (ROS) and defense secondary metabolites, respectively. While the OS from M. loreyi contained large amounts of previously reported insect elicitors, fatty acid-amino acid conjugates (FACs), the elicitor-active P. guttata’s OS contained no detectable FACs. Subsequently, elicitor activity associated with the high molecular mass fraction in OS of both herbivores was identified, and shown to promote ROS and metabolite accumulations in rice cells. Notably, the application of N-linolenoyl-Gln (FAC) alone had only negligible elicitor activity in rice cells; however, the activity of isolated elicitor fraction was substantially promoted by this FAC. Our results reveal that plants integrate various independent signals associated with their insect attackers to modulate their defense responses and reach maximal fitness in nature.

Constitutive and induced resistance in soybean interact to affect the performance of a herbivore and its parasitoid

This study examined how the induced defenses of soybean cultivars that vary in levels of constitutive resistance affect the developmental performance of the tobacco cutworm, Spodoptera litura, and its parasitoid, Meteorus pulchricornis. Both soybean cultivars were injured to induce defenses by either the beet armyworm, Spodoptera exigua, or by the aphid Aphis glycines. Both types of feeding on soybean plants had a negative effect on S. litura development, extending development time, reducing larva-to-pupa survival and decreasing adult body size. The impact of induction treatments increased linearly with increasing levels of constitutive resistance. Induction by the chewing herbivore was stronger than induction by the sucking herbivore in negatively affecting developmental parameters at both low and medium levels of soybean constitutive resistance, but had similar impacts on S. litura survival and adult body weight at the high level of constitutive resistance. Induction by both chewing and sucking herbivores extended parasitoid development time and decreased survival and fecundity, with effects on both development time and fecundity increasing linearly with levels of constitutive resistance, but without any effect on offspring survival. Thus, induction by the two herbivore types differed in impact on parasitoid performance depending on the level of constitutive resistance expressed in the soybean cultivar. The chewing herbivore was more effective than the sucking herbivore at low and medium levels of constitutive resistance, but there were no differences in parasitoid performance at the high level of constitutive resistance. Overall, whereas both chewing and sucking herbivores had negative effects on the development of S. litura and its parasitoid, the chewing herbivore was more effective than the sucking herbivore at low and medium levels of constitutive resistance, but similar to it at the high level.

Mutualism exploitation: predatory drosophilid larvae sugar-trap ants and jeopardize facultative ant-plant mutualism.

An open question in the evolutionary ecology of ant-plant facultative mutualism is how other members of the associated community can affect the interaction to a point where reciprocal benefits are disrupted. While visiting Qualea grandiflora shrubs to collect sugary rewards at extrafloral nectaries, tropical savanna ants deter herbivores and reduce leaf damage. Here we show that larvae of the fly Rhinoleucophenga myrmecophaga, which develop on extrafloral nectaries, lure potentially mutualistic, nectar-feeding ants and prey on them. Foraging ants spend less time on fly-infested foliage. Field experiments showed that predation (or the threat of predation) on ants by fly larvae produces cascading effects through three trophic levels, resulting in fewer protective ants on leaves, increased numbers of chewing herbivores, and greater leaf damage. These results reveal an undocumented mode of mutualism exploitation by an opportunistic predator at a plant-provided food source, jeopardizing ant-derived protection services to the plant. Our study documents a rather unusual case of predation of adult ants by a dipteran species and demonstrates a top-down trophic cascade within a generalized ant-plant mutualism.

Biosynthesis of 8-O-Methylated Benzoxazinoid Defense Compounds in Maize.

Benzoxazinoids are important defense compounds in grasses. Here, we investigated the biosynthesis and biological roles of the 8-O-methylated benzoxazinoids, DIM2BOA-Glc and HDM2BOA-Glc. Using quantitative trait locus mapping and heterologous expression, we identified a 2-oxoglutarate-dependent dioxygenase (BX13) that catalyzes the conversion of DIMBOA-Glc into a new benzoxazinoid intermediate (TRIMBOA-Glc) by an uncommon reaction involving a hydroxylation and a likely ortho-rearrangement of a methoxy group. TRIMBOA-Glc is then converted to DIM2BOA-Glc by a previously described O-methyltransferase BX7. Furthermore, we identified an O-methyltransferase (BX14) that converts DIM2BOA-Glc to HDM2BOA-Glc. The role of these enzymes in vivo was demonstrated by characterizing recombinant inbred lines, including Oh43, which has a point mutation in the start codon of Bx13 and lacks both DIM2BOA-Glc and HDM2BOA-Glc, and Il14H, which has an inactive Bx14 allele and lacks HDM2BOA-Glc in leaves. Experiments with near-isogenic maize lines derived from crosses between B73 and Oh43 revealed that the absence of DIM2BOA-Glc and HDM2BOA-Glc does not alter the constitutive accumulation or deglucosylation of other benzoxazinoids. The growth of various chewing herbivores was not significantly affected by the absence of BX13-dependent metabolites, while aphid performance increased, suggesting that DIM2BOA-Glc and/or HDM2BOA-Glc provide specific protection against phloem feeding insects.

Molecular evidence for biochemical diversification of phenolamide biosynthesis in rice plants.

Two phenolamides (PAs), p-coumaroylputrescine and feruloylputrescine strongly accumulate in rice (Oryza sativa cv. Nipponbare) leaves subjected to attack of chewing and sucking herbivores. Here we identified and characterized in vitro three novel rice genes that mediated coumaroyl-CoA/feruloyl-CoA conjugation to polyamines, putrescine and agmatine. Interestingly, two genes were highly specific for their polyamine substrates, encoding putrescine N-hydroxycinnamoyltransferase and agmatine N-hydroxycinnamoyltransferase, while the third enzyme could use both polyamines and it was therefore annotated as putrescine/agmatine N-hydroxycinnamoyltransferase. All genes were preferentially expressed in rice roots and developing flowers, and in addition, the putrescine/agmatine N-hydroxycinnamoyltransferase transcripts were strongly induced by wounding in the young rice leaves. Because the wound response of this gene was only partially suppressed in the jasmonoyl-L-isoleucine deficient plants (Osjar1), it suggests that its upregulation (as well as inducible PAs in rice) may be largely independent of jasmonoyl-L-isoleucine signaling pathway. The finding of three closely related genes with a similar and/or overlapping activity in PA biosynthesis provides another striking example of rapid diversification of plant metabolism in response to environmental stresses in nature.

Olfactory response of four aphidophagous insects to aphid- and caterpillar-induced plant volatiles

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.

Community structure and abundance of insects in response to early‐season aphid infestation in wild cabbage populations

1. Changes in the arthropod community structure can be attributed to differences in constitutively expressed plant traits or those that change depending on environmental conditions such as herbivory. Early‐season herbivory may have community‐wide effects on successive insect colonisation of host plants and the identity of the initially inducing insect may determine the direction and strength of the effects on the dynamics and composition of the associated insect community.2. Previous studies have addressed the effect of early infestation with a chewing herbivore. In the present study, the effect of early infestation was investigated with a phloem‐feeding aphid [Brevicoryne brassicae L. (Hemiptera, Aphididae)] on the insect community associated with three wild cabbage (Brassica oleracea L.) populations, which are known to differ in defence chemistry, throughout the season in field experiments.3. Aphid infestation had asymmetric effects on the associated insect community and only influenced the abundance of the natural enemies of aphids, but not that of chewing herbivores and their natural enemies. The effect size of aphid infestation further depended on the cabbage population.4. Aphid feeding has been previously reported to promote host‐plant quality for chewing herbivores, which has been attributed to antagonism between the two major defence signalling pathways controlled by the hormones salicylic acid (SA) and jasmonic acid (JA), respectively. Our results show no effects of early infestation by aphids on chewing herbivores, suggesting the absence of long‐term JA–SA antagonism.5. Investigating the effects of the identity of an early‐season coloniser and genotypic variation among plant populations on insect community dynamics are important in understanding insect–plant community ecology.

How plants handle multiple stresses: hormonal interactions underlying responses to abiotic stress and insect herbivory.

Adaptive plant responses to specific abiotic stresses or biotic agents are fine-tuned by a network of hormonal signaling cascades, including abscisic acid (ABA), ethylene, jasmonic acid (JA) and salicylic acid. Moreover, hormonal cross-talk modulates plant responses to abiotic stresses and defenses against insect herbivores when they occur simultaneously. How such interactions affect plant responses under multiple stresses, however, is less understood, even though this may frequently occur in natural environments. Here, we review our current knowledge on how hormonal signaling regulates abiotic stress responses and defenses against insects, and discuss the few recent studies that attempted to dissect hormonal interactions occurring under simultaneous abiotic stress and herbivory. Based on this we hypothesize that drought stress enhances insect resistance due to synergistic interactions between JA and ABA signaling. Responses to flooding or waterlogging involve ethylene signaling, which likely reduces plant resistance to chewing herbivores due to its negative cross-talk with JA. However, the outcome of interactions between biotic and abiotic stress signaling is often plant and/or insect species-dependent and cannot simply be predicted based on general knowledge on the involvement of signaling pathways in single stress responses. More experimental data on non-model plant and insect species are needed to reveal general patterns and better understand the molecular mechanisms allowing plants to optimize their responses in complex environments.

OsWRKY53, a versatile switch in regulating herbivore-induced defense responses in rice

ABSTRACTWRKY proteins, which belong to a large family of plant-specific transcription factors, play important roles in plant defenses against pathogens and herbivores by regulating defense-related signaling pathways. Recently, a rice WRKY transcription factor OsWRKY53 has been reported to function as a negative feedback modulator of OsMPK3/OsMPK6 and thereby to control the size of the investment a rice plant makes to defend against a chewing herbivore, the striped stem borer Chilo suppressalis. We investigated the performance of a piecing-sucking herbivore, the brown planthopper (BPH) Nilaparvata lugens, on transgenic plants that silence or overexpress OsWRKY53, and found that OsWRKY53 activates rice defenses against BPH by activating an H2O2 burst and suppressing ethylene biosynthesis. These findings suggest that OsWRKY53 functions not only as a regulator of plants' investment in specific defenses, but also as a switch to initiate new defenses against other stresses, highlighting the versatility and importance of OsWRKY53 in herbivore-induced plant defenses.

A conserved pattern in plant-mediated interactions between herbivores.

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.

Ecology of some lesser-studied introduced ant species in Hawaiian forests

Invasive ants can have strong ecological effects on native arthropods, but most information on this topic comes from studies of a handful of ant species. The ecological impacts of the many additional introduced ‘tramp’ ant species are largely unknown. In mesic upland forests of O‘ahu, Hawai‘i, ten species of introduced ants were found on four species of understory trees. However, these ants were generally uncommon and occurred at relatively low densities in this habitat type. The most common and abundant ant was Plagiolepis alluaudi, which favored the native tree Pipturus albidus. Ecological effects of ants on arthropods were found to be modest, with overall arthropod community composition not significantly different between ant-occupied and ant-free trees. Most taxonomic groups were similar in abundance and richness between ant-occupied and ant-free trees, except adventive Coleoptera, adventive Hemiptera, and Lepidoptera were less abundant on ant-occupied trees, and adventive Coleoptera and adventive Hemiptera also had lower richness on ant-occupied trees. Among Lepidoptera, caterpillars of two endemic groups had significantly lower incidences on trees with higher ant abundances, while other caterpillars did not. Arthropod trophic structure was largely unaffected, except that chewing herbivores comprised a smaller fraction of biomass on ant-occupied trees. While overall ecological effects were weak in comparison to prior results involving other ant species in Hawai‘i, some of the impacts suggest that higher densities of these introduced ant species could result in similar interactions with arthropods as those of the better-studied invasive ant species.

Change in herbivore insect communities from adjacent habitats in a transitional region

Ecotonal areas between habitats vary in floristic composition and phenological patterns of the vegetation. In these regions, the habitat structure and plant phenology potentially affect the diversity of associated herbivore insects guilds (chewing and sap-sucking). In this study, the following hypotheses were tested in three habitats (cerrado sensu strictu, dry forest and riparian forest) in a Cerrado–Caatinga transitional area: (1) There is a variation in composition, richness and abundance of herbivore insects associated with the canopy among adjacent habitats; (2) habitats with higher variation in canopy cover support lower richness and abundance of herbivore insects; (3) habitats with more tree species and higher tree density support higher richness and abundance of herbivore insects; and (4) differences in species composition of herbivore insects among adjacent habitats are the main source of diversity in transitional habitats. The community of herbivore insects differed among the habitats, being similar between riparian and dry forests and different from cerrado. Canopy cover variation, richness and density of trees did not influence richness and abundance of chewing herbivores. A negative effect of canopy cover variation was observed in richness and abundance of sap-sucking herbivore insects, and a negative effect of tree density was observed on abundance of sap-sucking herbivore insects. A higher importance of beta diversity among habitats was observed if compared to beta diversity within habitats (54 %). This study points out that strategies of conservation must consider the entire vegetation mosaics to protect a higher number of species, and to keep these regions with high diversity.

Reexamination of Chlorophyllase Function Implies Its Involvement in Defense against Chewing Herbivores

Chlorophyllase (CLH) is a common plant enzyme that catalyzes the hydrolysis of chlorophyll to form chlorophyllide, a more hydrophilic derivative. For more than a century, the biological role of CLH has been controversial, although this enzyme has been often considered to catalyze chlorophyll catabolism during stress-induced chlorophyll breakdown. In this study, we found that the absence of CLH does not affect chlorophyll breakdown in intact leaf tissue in the absence or the presence of methyl-jasmonate, which is known to enhance stress-induced chlorophyll breakdown. Fractionation of cellular membranes shows that Arabidopsis (Arabidopsis thaliana) CLH is located in the endoplasmic reticulum and the tonoplast of intact plant cells. These results indicate that CLH is not involved in endogenous chlorophyll catabolism. Instead, we found that CLH promotes chlorophyllide formation upon disruption of leaf cells, or when it is artificially mistargeted to the chloroplast. These results indicate that CLH is responsible for chlorophyllide formation after the collapse of cells, which led us to hypothesize that chlorophyllide formation might be a process of defense against chewing herbivores. We found that Arabidopsis leaves with genetically enhanced CLH activity exhibit toxicity when fed to Spodoptera litura larvae, an insect herbivore. In addition, purified chlorophyllide partially suppresses the growth of the larvae. Taken together, these results support the presence of a unique binary defense system against insect herbivores involving chlorophyll and CLH. Potential mechanisms of chlorophyllide action for defense are discussed.

Additive and non‐additive effects of birch genotypic diversity on arthropod herbivory in a long‐term field experiment

Herbivores are important drivers of plant population dynamics and community composition in natural and managed systems. Intraspecific genetic diversity of long‐lived plants like trees might shape patterns of herbivory by different guilds of herbivores that trees experience through time. However, previous studies on plant genetic diversity effects on herbivores have been largely short‐term. We investigated how tree genotypic variation and diversity influence herbivory of silver birchBetula pendulain a long‐term field experiment. Using clones of eight genotypes, we constructed experimental plots consisting of one, two, four or eight genotypes, and measured damage by five guilds of arthropod herbivores twice a year over three different years (four, six and nine years after the experiment was established). Genotypes varied significantly for most types of herbivore damage, but genotype resistance rankings often shifted over time, and none of the clones was more resistant than all others to all types of herbivores. At the plot level, birch genotypic diversity had significant positive additive effect on leaf rollers and negative non‐additive effects on chewing herbivores and gall makers. In contrast, leaf‐mining and leaf‐tying damage was not influenced by birch genotypic diversity. Within diverse plots, the direction of genotypic diversity effects varied depending on birch genotype, some having lower and some having higher herbivory in mixed stands. This research highlights the importance of long‐term studies including different feeding guilds of herbivores to understand the effects of plant genetic diversity on arthropod communities. Different responses of various feeding guilds to genotypic diversity and shifts in resistance of individual genotypes over time indicate that genotypic mixtures are unlikely to result in overall reduction in herbivory over time.

Transcriptional responses of Arabidopsis thaliana to chewing and sucking insect herbivores.

We tested the hypothesis that Arabidopsis can recognize and respond differentially to insect species at the transcriptional level using a genome wide microarray. Transcriptional reprogramming was characterized using co-expression analysis in damaged and undamaged leaves at two times in response to mechanical wounding and four insect species. In all, 2778 (10.6%) of annotated genes on the array were differentially expressed in at least one treatment. Responses differed mainly between aphid and caterpillar and sampling times. Responses to aphids and caterpillars shared only 10% of up-regulated and 8% of down-regulated genes. Responses to two caterpillars shared 21 and 12% of up- and down-regulated genes, whereas responses to the two aphids shared only 7 and 4% of up-regulated and down-regulated genes. Overlap in genes expressed between 6 and 24 h was 3–15%, and depended on the insect species. Responses in attacked and unattacked leaves differed at 6 h but converged by 24 h. Genes responding to the insects are also responsive to many stressors and included primary metabolism. Aphids down-regulated amino acid catabolism; caterpillars stimulated production of amino acids involved in glucosinolate synthesis. Co-expression analysis revealed 17 response networks. Transcription factors were a major portion of differentially expressed genes throughout and responsive genes shared most of the known or postulated binding sites. However, cis-element composition of genes down regulated by the aphid M. persicae was unique, as were those of genes down-regulated by caterpillars. As many as 20 cis-elements were over-represented in one or more treatments, including some from well-characterized classes and others as yet uncharacterized. We suggest that transcriptional changes elicited by wounding and insects are heavily influenced by transcription factors and involve both enrichment of a common set of cis-elements and a unique enrichment of a few cis-elements in responding genes.

Bison grazing increases arthropod abundance and diversity in a tallgrass prairie.

How grazing-induced ecosystem changes by ungulates indirectly affect other consumers is a question of great interest. I investigated the effect of grazing by American Bison (Bos bison L.) on an arthropod community in tallgrass prairie. Grazing increased the abundance of arthropods, an increase that was present in both herbivorous and carnivorous assemblages, but not in detritivores. The increase in herbivores and reduction in plant biomass from grazing resulted in an arthropod herbivore load almost three times higher in grazed plots compared with controls. Among herbivores, the sap-feeding insect guild was dramatically more abundant, while chewing herbivores were not affected. Herbivorous and carnivorous arthropod richness was higher in grazed plots, although the response was strongest among herbivores. Arthropod abundance on individual grasses and forbs was significantly higher in grazed areas, while plant type had no effect on abundance, indicating that the change was ecosystem-wide and not simply in response to a reduction in grass biomass from grazing. The response of arthropods to grazing was strongest in the early part of the growing season. Published research shows that ungulate grazing, although decreasing available biomass to other consumers, enhances plant quality by increasing nitrogen level in plants. The arthropod results of this study suggest higher plant quality outweighs the potential negative competitive effects of plant biomass removal, although other activities of bison could not be ruled out as the causative mechanism. Because arthropods are extremely abundant organisms in grasslands and a food source for other consumers, bison may represent valuable management tools for maintaining biodiversity.