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Grasses accumulate large amounts of silicon (Si) which acts as a highly effective physical defence against insect herbivory, however recent evidence shows that Si supplementation also modifies plant secondary metabolite concetrations. Changes in plant secondary metabolites concentrations can have cascading effects on higher trophic levels, such as parasitoids, as they are dependent on the host herbivore for growth and development. However, relatively little is known about how Si application affects higher trophic levels. We examined the effects of Si addition on alkaloid content in leaves of Phalaris aquatica (Poaceae) and the effect on interactions between an aphid (Rhopalosiphum padi) and its parasitoid (Aphidius colemani). Si supplementation had no effect on aphid abundance or parasitism rate. Adult aphids, aphid mummies (parasitised aphids) and the emergent parasitoids were, however, significantly smaller on Si+ plants. Parasitoid traits (size and emergence) were correlated with aphid mummy size. Si addition reduced parasitoid emergence rate and size due to reduced host mummy size, in addition, significantly fewer females emerged from mummies on Si+ plants. Aphid infestation significantly altered alkaloids concentrations, reducing gramine by 80% while increasing tryptamine by 91% in Si- plants. Si addition reduced aphid-induced tryptamine concentrations by 64% and increased 5-MeO-tryptamine by over 800% in control and 142% in aphid infested plants. Our results show that while Si addition has modest impacts on the herbivore, it significantly alters secondary metabolites and has stronger effects on the higher trophic level through changes in the quality of the parasitised host.

Silicon (Si) has been widely reported to improve plant resistance to water stress via various mechanisms including cuticular Si deposition to reduce leaf transpiration. However, there is limited understanding of the effects of Si on stomatal physiology, including the underlying mechanisms and implications for resistance to water stress. We grew tall fescue (Festuca arundinacea Schreb. cv. Fortuna) hydroponically, with or without Si, and treated half of the plants with 20% polyethylene glycol to impose physiological drought (osmotic stress). Scanning electron microscopy in conjunction with X-ray mapping found that Si was deposited on stomatal guard cells and as a sub-cuticular layer in Si-treated plants. Plants grown in Si had a 28% reduction in stomatal conductance and a 23% reduction in cuticular conductance. When abscisic acid was applied exogenously to epidermal leaf peels to promote stomatal closure, Si plants had 19% lower stomatal aperture compared to control plants (i.e. increased stomatal sensitivity) and an increased efflux of guard cell K+ ions. However, the changes in stomatal physiology with Si were not substantial enough to improve water stress resistance, as shown by a lack of significant effect of Si on water potential, growth, photosynthesis and water-use efficiency. Our findings suggest a novel underlying mechanism for reduced stomatal conductance with Si application; specifically, that Si deposition on stomatal guard cells promotes greater stomatal sensitivity as mediated by guard cell K+ efflux.

The role of silicon (Si) in alleviating the effects of biotic and abiotic stresses, including defence against insect herbivores, in plants is widely reported. Si defence against insect herbivores is overwhelmingly studied in grasses (especially the cereals), many of which are hyper-accumulators of Si. Despite being neglected, legumes such as soybean (Glycine max) have the capacity to control Si accumulation and benefit from increased Si supply. We tested how Si supplementation via potassium, sodium or calcium silicate affected a soybean pest, the native budworm Helicoverpa punctigera Wallengren (Lepidoptera: Noctuidae). Herbivory reduced leaf biomass similarly in Si-supplemented (+Si) and non-supplemented (–Si) plants (c. 29 and 23%, respectively) relative to herbivore-free plants. Both Si supplementation and herbivory increased leaf Si concentrations. In relative terms, herbivores induced Si uptake by c. 19% in both +Si and –Si plants. All Si treatments reduced H. punctigera relative growth rates (RGR) to a similar extent for potassium (−41%), sodium (−49%) and calcium (−48%) silicate. Moreover, there was a strong negative correlation between Si accumulation in leaves and herbivore RGR. To our knowledge, this is only the second report of Si-based herbivore defence in soybean; the rapid increase in leaf Si following herbivory being indicative of an induced defence. Taken together with the other benefits of Si supplementation of legumes, Si could prove an effective herbivore defence in legumes as well as grasses.

Grasses accumulate large concentrations of silicon (Si) which alleviates a range of stresses including defence against herbivores. Likewise, grasses symbiotically associate with foliar Epichloë-fungal endophytes which provide herbivore defence, mainly via the production of alkaloids. Some Epichloë-endophytes increase foliar Si concentrations, particularly in tall fescue (Festuca arundinacea) but also in perennial ryegrass (Lolium perenne); it is unknown whether this impacts herbivores. Likewise, while Si is primarily a physical defence against herbivores, it can also affect defensive secondary metabolites; Si supply might therefore also affect alkaloids produced by Epichloë-endophytes, however, this remains untested. We grew tall fescue and perennial ryegrass in a factorial combination with or without Si supplementation, in the absence or presence of a chewing herbivore; Helicoverpa armigera. Grasses were associated with four different Epichloë-endophyte strains (tall fescue: AR584; perennial ryegrass: AR37, AR1, or wild type) or as Epichloë-free controls. Specifically, we assessed how Si supply and Epichloë-endophyte presence impacts plant growth and chemistry, and how their interaction with herbivory affects foliar Si concentrations and alkaloid production. Subsequently, their effects on H. armigera relative growth rates (RGR) were evaluated. In Fescue, the AR584-endophyte increased constitutive (herbivore-free) and induced (herbivore-inoculated) silicon concentrations when Si was supplied. In perennial ryegrass, AR37-endophyte increased constitutive and induced silicon concentration, meanwhile, AR1-endophyte increased constitutive levels only. Si supply and herbivory did not affect alkaloids produced by AR584- or AR1/Wt-endophyte in tall fescue and perennial ryegrass, respectively. However, Si suppressed herbivore-induced production of alkaloids in the AR37-endophyte perennial ryegrass association. Si was a more effective defence in tall fescue than perennial ryegrass, significantly reducing H. armigera RGR. Our results suggest that Si reduced herbivore performance to such an extent in tall fescue that it was operating at maximum effect and endophyte-mediated increases in Si concentration made no further difference. Si had a more modest impact on herbivores in perennial ryegrass, potentially linked to silicon decreasing herbivore feeding and thus, suppressing herbivore-induced alkaloids. We provide novel evidence that increased Si concentrations in some cases interact with endophyte-produced chemical defences, which could ultimately impact plant resistance to herbivores.

1) Crop loss due to insect herbivory is one of the largest challenges facing the agricultural industry. As herbivore populations continue to grow in light of global change, securing crop resources is becoming increasingly critical. Silicon (Si) has been shown to effectively mitigate the adverse effects of herbivores such as the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae), in crop species (namely grasses), that have evolved the ability to uptake large amounts of Si through their roots and accumulate it in aboveground tissues. Nevertheless, the effectiveness of Si accumulation as a plant defence against herbivory in the short term, and its consequential effects on alternative defence responses, remain unclear.2) We conducted two discrete experiments to determine the short-term dynamics of Si, chemical defences and resistance to herbivory in the model grass, Brachypodium distachyon: 1) Both Si-supplemented (+Si) and control (-Si) plants were treated with methyl jasmonate (MeJA) as a form of simulated herbivory and we measured the interplay of Si accumulation, the phytohormones jasmonic acid (JA) and salicylic acid (SA), and carbon-based defences over 24 hr. 2) We exposed H. armigera larvae to B. distachyon plants grown under three conditions: +Si, -Si, or treated with Si only once H. armigera feeding began. We measured the effect of short-term plant exposure to Si on H. armigera performance and plant resistance.3) MeJA-induced Si accumulation occurred as early as 6 hr after treatment via increased JA concentrations. Si supplementation decreased SA concentrations, which could have implications on additional downstream defences. We show a trade-off between Si and phenolics in untreated plants, but this relationship was weakened upon MeJA treatment. Although foliar Si concentrations remained lower, within 72 hr of exposure to Si, plants obtained virtually the same level of resistance to H. armigera as plants exposed to Si for over 30 days. H. armigera feeding also accelerated Si deposition after 6 hr of exposure to Si, however, in as little as 24 hr, levels of Si deposition were similar to plants exposed to Si long term.4) In addition to its well-documented role as a long-term defence against herbivores, we demonstrate that, over short-term temporal scales, Si accumulation responds to herbivore signals and impacts on plant defence machinery. Further, we provide novel evidence that plants can rapidly incorporate Si into their tissues to mitigate the adverse effects of herbivory as effectively as plants exposed to Si long term.

Plants deploy an arsenal of chemical and physical defenses against arthropod herbivores, but it may be most cost efficient to produce these only when attacked. Herbivory activates complex signaling pathways involving several phytohormones, including jasmonic acid (JA), which regulate production of defensive compounds. The Poaceae also have the capacity to take up large amounts of silicon (Si), which accumulates in plant tissues. Si accumulation has antiherbivore properties, but it is poorly understood how Si defenses relate to defense hormone signaling. Here we show that Si enrichment causes the model grass Brachypodium distachyon to show lower levels of JA induction when attacked by chewing herbivores. Triggering this hormone even at lower concentrations, however, prompts Si uptake and physical defenses (e.g., leaf hairs), which negatively impact chewing herbivores. Removal of leaf hairs restored performance. Crucially, activation of such Si-based defense is herbivore-specific and occurred only in response to chewing and not fluid-feeding (aphid) herbivores. This aligned with our meta-analysis of 88 studies that showed Si defenses were more effective against chewing herbivores than fluid feeders. Our results suggest integration between herbivore defenses in a model Si-accumulating plant, which potentially allows it to avoid unnecessary activation of other costly defenses.

Changes in insect herbivore performance under elevated atmosphere carbon dioxide concentrations e[CO2] are often driven by changes in the nutritional and defensive chemistry of their host plants. Studies addressing how the prolific pest cotton bollworm (Helicoverpa armigera) responds to e[CO2] show that performance usually declines, often associated with lower nutritional (e.g. nitrogen (N) concentrations) quality of host plants under e[CO2]. We investigated the impacts of e[CO2] on nutritional quality and anti-herbivore (jasmonate) defensive signalling in lucerne (Medicago sativa) when challenged by H. armigera. While foliar N decreased under e[CO2], other aspects of nutritional quality (soluble protein, amino acids, foliar C:N) were largely unaffected, potentially due to increased root nodulation under e[CO2]. In contrast, e[CO2] greatly reduced jasmonate signalling in M. sativa following H. armigera attack; jasmonic acid concentrations were ca. 56% lower in attacked plants grown under e[CO2]. Concurrent with this, relative growth rates of H. armigera were ca. 66% higher when feeding on e[CO2]-grown plants. In contrast with previous reports, which we meta-analytically summarise, we provide the first evidence that H. armigera performance can increase under e[CO2]. This may occur in plants, such as M. sativa, where e[CO2] has limited impacts on nutritional quality yet reduces jasmonate defence signalling.

Grasses accumulate large amounts of silicon (Si) which acts as a highly effective physical defence against insect herbivory, however recent evidence shows that Si supplementation also modifies plant secondary metabolite concetrations. Changes in plant secondary metabolites concentrations can have cascading effects on higher trophic levels, such as parasitoids, as they are dependent on the host herbivore for growth and development. However, relatively little is known about how Si application affects higher trophic levels. We examined the effects of Si addition on alkaloid content in leaves of Phalaris aquatica (Poaceae) and the effect on interactions between an aphid (Rhopalosiphum padi) and its parasitoid (Aphidius colemani). Si supplementation had no effect on aphid abundance or parasitism rate. Adult aphids, aphid mummies (parasitised aphids) and the emergent parasitoids were, however, significantly smaller on Si+ plants. Parasitoid traits (size and emergence) were correlated with aphid mummy size. Si addition reduced parasitoid emergence rate and size due to reduced host mummy size, in addition, significantly fewer females emerged from mummies on Si+ plants. Aphid infestation significantly altered alkaloids concentrations, reducing gramine by 80% while increasing tryptamine by 91% in Si- plants. Si addition reduced aphid-induced tryptamine concentrations by 64% and increased 5-MeO-tryptamine by over 800% in control and 142% in aphid infested plants. Our results show that while Si addition has modest impacts on the herbivore, it significantly alters secondary metabolites and has stronger effects on the higher trophic level through changes in the quality of the parasitised host.

Prior feeding by insect herbivores frequently affects plant quality for herbivores that subsequently feed on the plant. Facilitation occurs when one herbivore improves plant quality for other herbivores, including when the former compromises plant defenses. Silicon (Si) is an important defense in grasses that increases following activation of the jasmonic acid (JA) pathway. Given that aphids often stimulate the salicylic acid (SA) pathway, we hypothesized that this could reduce Si defense because of the well documented antagonistic cross-talk between SA and JA. We tested this in the model grass Brachypodium distachyon with and without Si (+Si and &minus, Si, respectively), half of the plants were exposed to aphids (Rhopalosiphum padi) and half remained aphid-free. Aphid-free and aphid-exposed plants were then fed to chewing herbivores (Helicoverpa armigera). Aphids triggered higher SA concentrations which suppressed JA concentrations but this did not affect foliar Si. Chewing herbivores triggered higher JA concentrations and induced Si uptake, regardless of previous feeding by aphids. Chewer growth rates were not impacted by prior aphid herbivory but were reduced by 75% when feeding on +Si plants. We concluded that aphids caused phytohormonal cross-talk but this was overridden by chewing herbivory that also induced Si uptake.

Grasses accumulate large amounts of silicon (Si) which is deposited in trichomes, specialised silica cells and cell walls. This may increase leaf toughness and reduce cell rupture, palatability and digestion. Few studies have measured leaf mechanical traits in response to Si, thus the effect of Si on herbivores can be difficult to disentangle from Si-induced changes in leaf surface morphology. We assessed the effects of Si on Brachypodium distachyon mechanical traits (specific leaf area (SLA), thickness, leaf dry matter content (LDMC), relative electrolyte leakage (REL)) and leaf surface morphology (macrohairs, prickle, silica and epidermal cells) and determined the effects of Si on the growth of two generalist insect herbivores (Helicoverpa armigera and Acheta domesticus). Si had no effect on leaf mechanical traits, however, Si changed leaf surface morphology: silica and prickle cells were on average 127% and 36% larger in Si supplemented plants, respectively. Prickle cell density was significantly reduced by Si, while macrohair density remained unchanged. Caterpillars were more negatively affected by Si compared to crickets, possibly due to the latter having a thicker and thus more protective gut lining. Our data show that Si acts as a direct defence against leaf-chewing insects by changing the morphology of specialised defence structures without altering leaf mechanical traits.

Galling insects have the unique ability to control the secondary chemistry of their host plants. Galled plant tissue often has higher concentrations of these secondary compounds compared with surrounding plant material. In this study, we used nuclear magnetic resonance (NMR) spectroscopy metabolomics techniques to investigate the effect of galls induced by Psephodiplosis rubi Kolesik, on the profiles of secondary metabolites in its two Rubus host plants in subtropical rainforest. We isolated four triterpenoid compounds from both Rubus species. Concentrations of individual triterpenes were not significantly different in galled and non‐galled leaves; however, the concentration of 2‐oxopolomic acid was significantly negatively correlated with gall abundance. At the interspecific level, non‐galled leaves from both Rubus species were chemically indistinguishable in principal component analysis analyses, while galled leaves showed slight chemical differences. Surprisingly, there was very little difference in metabolic profiles of galled and non‐galled leaves at the intraspecific level. Using NMR, we were able to detect slightly different chemical responses in two congeneric host plant species induced by P. rubi galls. We demonstrate how NMR can be used to detect interspecific chemical differences of plants in ecological studies.

Solanum inaequilaterum is a perennial shrub, endemic to the evergreen subtropical rainforest of southern Queensland and northern New South Wales, Australia. An undescribed species of gall midge (Diptera: Cecidomyiidae) was found to cause numerous round hairy galls on leaves, leaf stalks and stems. The new species is described and named Dasineura inaequilaterae Kolesik sp. nov. It is the first Dasineura known to feed on a host plant from the family Solanaceae worldwide.

An undescribed species of gall midge causing hairy spheroidal leaf galls on Rubus nebulosus and Rubus moorei (Rosaceae) in subtropical Australia was investigated in a study on the dynamics of parasitoids of gall-inducing insects. Here, the gall midge is described and named Psephodiplosis rubi Kolesik, sp. nov. in a new genus Psephodiplosis Kolesik, gen. nov. (supertribe Cecidomyiidi) erected to contain it. The new species is the first described gall midge that induces galls on Rosaceae in Australia. Similar galls have been found previously on Rubus moluccanus in Indonesia and on Rubus alceifolius in Thailand, so it is possible that the geographical range of Psephodiplosis rubi includes not only Australia, but also countries of South East Asia.

The idea that galling insects actively manipulate host plant chemistry has been previously documented but has not been quantified across a range of galler and host plant taxa. We present the first quantitative review of the relationship between insect galling and levels of secondary metabolites in host plants. Using meta-analytic techniques, we examined this relationship across 40 galler and host plant species combinations. We found that galling insects are associated with significantly higher levels of tannins and phenolics; however, no difference was found for volatiles. Hymenoptera, Diptera and Hemiptera were associated with higher levels of secondary metabolites; however, only Hymenoptera was significant. The climatic zone of the study area did not explain significant differences in gall-induced secondary metabolites. Overall the results show that the ability of galling insects to manipulate host plant secondary chemistry is widespread across insect and plant taxa. The evolutionary success of galling insects may be in part due to this unique ability.

International audience; Thermal requirements for fl ight in butterfl ies is determined by a combination of external factors, behaviour and physical constraints. Thorax temperature of 152 butterfl ies was monitored with an infra-red thermometer in controlled laboratory conditions. The temperature at take-off varied from 13.4 degrees C, for a female Heteronympha merope to 46.3 degrees C, for a female Junonia villida. Heteronympha merope, an understorey species, had the lowest recorded take-off temperatures, with females fl ying at a much lower thorax temperatures than males. Among the tested butterfl y species, warming-up rate was positively correlated with take-off temperature and negatively with body mass. Wing loading is a major variable in determining the thorax fl ight temperature. Butterfl ies with the highest wing-loadings experienced the highest thorax temperatures at take-off. A notable exception to this rule is Trapezites symmomus, the only Hesperiidae of our data set, which had thorax fl ight temperatures of 31.5 degrees C and 34.5 degrees C, well within the range of the observed butterfl ies, despite a wing load ca. fi ve times higher. The high thorax temperature recorded in J. villida is probably linked to its high fl ight speed. The results highlight the importance of physical constraints such as body size on the thermal requirements for fl ight across a range of butterfl y species.

Parasitoids play an important role in ecosystem functioning through their influence on herbivorous insect populations. Theoretical and experimental evidence suggest that increased species richness can enhance and stabilize ecosystem function. It is important to understand how richness-driven functional relationships change across environmental gradients. We investigated how temperature affected the relationship between parasitoid richness and parasitism rate in a guild of gall-parasitoids along an elevational gradient. We collected galls at 15 sites along five elevational gradients (between 762 m and 1145 m asl) on six occasions over a year. A total of 1902 insects, including 1593 parasitoids, were reared from 12 402 galls. Parasitism rate increased significantly with temperature on all sampling occasions, except December and February. We found a significant, positive richness–parasitism relationship. This relationship, however, was weaker at higher elevations which may be linked to decreased functional efficiency of parasitoids at lower temperatures. Temporal variability in parasitism rate and parasitoid richness were significantly related, regardless of temperature. A stable functional guild of this kind may provide a more reliable ecosystem service under environmental changes.

The genus Flindersia (Rutaceae) comprises 17 species of mostly Australian endemic trees. Although most species are restricted to rainforests, four have evolved to grow in semi-arid and arid environments. In this study, the leaf alkaloid diversity of rainforest and semi-arid/arid zone adapted Australian Flindersia were compared by LC/MS-MS and NMR spectroscopy. Contrary to expectations, Flindersia alkaloid diversity was strongly correlated with environmental aridity, where species predominating in drier regions produced more alkaloids than their wet rainforest congenerics. Rainforest species were also more chemically similar to each other than were the four semi-arid/arid zone species. There was a significant relationship between the presence of alkaloid structural classes and phylogenetic distance, suggesting that alkaloid profiles are influenced by both genetic and environmental factors. The results suggest that the radiation of Flindersia species out of the rainforest and into drier environments has promoted the evolution of unique alkaloid diversity. Plants growing in arid and semi-arid regions of Australia may represent an untapped source of undescribed specialised metabolites.

Patterns of species assemblages along altitudinal gradients provide insights into ecological patterns and processes. Parasitoids play an important role in ecosystems, yet there are few studies on parasitoid assemblages, partly because of difficulty in identification of parasitoids to species or their surrogates, morphospecies. The aim of this study is to assess differences in assemblages of parasitic Hymenoptera along an altitudinal gradient at both the family and morphospecies level, and the potential of environmental variables for explaining these differences. We sampled from four replicate sites at each of five elevations (approximately 300, 500, 700, 900 and 1100m above sea level (asl)) within continuous subtropical rainforest of the Border Ranges National Park in New South Wales, Australia. At each site, we operated a Malaise trap for 10 days in December 2011 and again in February 2012. Parasitic wasps were sorted to families, and two of the common families (Diapriidae and Ichneumonidae) were further sorted to morphospecies. At both family and morphospecies level, we found gradual altitudinal turnover with a clear separation between upland' (900 and 1100masl) and lowland' (300 and 500masl) assemblages. Unlike parallel studies of some other major taxa, we did not detect distinct assemblages at each altitude at the taxonomic resolution used. In addition, response patterns were different between the two sampling occasions. Changes in assemblage composition were best explained by either altitude or temperature. Although parasitic Hymenoptera do not respond as clearly to changes in altitude as some other taxa, the changes detected here, even at the family level, are important given the role parasitoids play in ecosystem functioning.