Undamaged Plants Research Articles

Pathways for plant‐mediated negative feedback to insect herbivores: accounting for non‐linear effects of larval density on plant quality and quantity

AbstractChanges in plant traits induced by herbivore damage can produce a negative feedback to increasing herbivore densities. Several aspects of these plant‐mediated feedbacks are predicted to influence herbivore population dynamics, but the degree to which feedbacks are non‐linear, whether their strength varies among plant genotypes, and whether they occur via changes in the quality of plant tissue or the amount of tissue available have rarely been examined. In this study, we damaged five genotypes of the perennial weed Carolina horsenettle, Solanum carolinense L. (Solanaceae), with eight densities of the false potato beetle, Leptinotarsa juncta (Germar) (Coleoptera: Chysomelidae, Chrysomelini). To account for plant quantity available to beetles, we measured the leaf area of each plant after imposing larval density treatments. We then measured the oviposition preference of L. juncta adults for damaged vs. undamaged plants and the relative growth rate (RGR) of L. juncta larvae on each plant, as well as plant trypsin proteinase inhibitor (tryPI) expression. We found that L. juncta females strongly preferred to oviposit on undamaged S. carolinense, and that preference for a plant decreased as the density of damaging larvae increased. In addition, we found decreased larval RGR and increased production of tryPIs with greater initial larval density. Effects of larval density on insect preference, performance, and tryPI expression were linear and did not vary among plant genotypes. Larval density effects were not solely due to reduced plant quantity, as plant leaf area had no effect on oviposition preference or larval performance. This suggests that effects were most likely due to changes in the quality of plant tissue. Thus, negative feedback on increasing larval density can be mediated by the effects of induced resistance on both oviposition preference and larval performance. Published models suggest that the linear, quality‐mediated feedbacks observed in this experiment are predicted to stabilize herbivore population dynamics.

Plant communication in a widespread goldenrod: keeping herbivores on the move

Summary Plant communication has been documented in over 35 plant species spanning 16 families to date; however, the underlying mechanisms through which it shapes plants' ecological interactions remain less clear. Using a combination of field/laboratory bioassays, headspace volatile and leaf chemical analyses in tall goldenrod (Solidago altissima), we tested the hypothesis that plant‐to‐plant communication affects the performance, feeding and movement behaviour of herbivores by changing plants' chemical phenotypes. We found that plant communication accelerates herbivore movement between host plants while simultaneously reducing herbivory. This suggests that plant communication can limit herbivore loads by keeping herbivores on the move between host plants. We demonstrate that volatile chemicals emitted from herbivore‐attacked plants are sufficient to explain metabolic responses in and ecological consequences for the exposed neighbour plant. Volatile profiles of beetle (Trirhabda virgata)‐damaged and undamaged plants show substantial compositional differences, especially in sesquiterpene emission, indicating that these differences have the potential to provide neighbouring plants with specific information about herbivores in the vicinity. Despite qualitative and phenological differences in plants' metabolic responses to herbivory‐induced volatile organic compounds (VOCs) and feeding herbivores, herbivores nonetheless respond similarly to directly damaged plants and plants exposed to VOCs from damaged neighbours as if they were of equivalently poor quality. This study suggests that by enlarging the spatial scale at which induced resistance affects the distribution of plant chemical phenotypes in plant populations, VOC‐mediated plant communication alters the movement behaviour and performance of herbivores. A lay summary is available for this article.

Differing Behavioural Responses of Bemisia tabaci MEAM1 and MED to Cabbage Damaged by Conspecifics and Heterospecifics

The whitefly Bemisia tabaci is a serious pest with an extensive host range. Previous research has shown that B. tabaci is a species complex with many cryptic species or biotypes and that the two most important species are MEAM1 (Middle East-Minor Asia 1) and MED (Mediterranean genetic group). MEAM1 and MED are known to differ in their preference for cabbage, Brassica oleracea, as a host plant, however, the mechanism underlying this preference is unknown. In the current study, a host choice experiment showed that MED prefers to settle and oviposit on undamaged cabbage plants rather than MED-damaged cabbage plants. However, MEAM1 prefers MED-damaged cabbage plants to undamaged plants and does not exhibit a significant preference for undamaged or MEAM1-damaged cabbage plants. On the basis of gas chromatography-mass spectrometry (GC-MS) analysis, the following volatiles were released in larger quantities from Q-damaged cabbage plants than from undamaged plants: 2-ethyl-1-hexanol, benzenemethanol, (E)-2-decenol, benzaldehyde, nonanal, acetic acid geraniol ester, 4-hydroxy-4-methyl-2-pentanone, decane, and α-longipinene. Only one volatile, 4-hydroxy-4-methyl-2-pentanone, was released in greater quantities from MEAM1-damaged cabbage plants than from undamaged plants. Our results suggest that differences in herbivore-induced host volatile release may help explain the differences between the preference of B. tabaci MEAM1 and MED for cabbage as a host.

Plant-plant interactions affect the susceptibility of plants to oviposition by pests but are disrupted by ozone pollution

Plant volatiles convey information about the physiological status of a plant to other organisms, including neighboring plants and herbivores searching for a suitable oviposition site. Tropospheric air pollutants, such as ozone, can impair plant–plant interactions, e.g. by degrading volatile signals. In this study, field and laboratory experiments were conducted to assess whether plant–plant interactions between damaged and undamaged cabbage (Brassica oleracea var. capitata) plants provide receivers with a competitive advantage by altering their susceptibility to oviposition by the specialist herbivores Pieris brassicae and Plutella xylostella. We tested the phenomenon in oviposition choice tests under ambient and enriched ozone conditions. We also investigated the effect of receiving a signal from a different Brassica oleracea cultivar group, in this study broccoli (Brassica oleracea var italica), on the susceptibility to oviposition of cabbage plants. Cabbage plants exposed to damaged cabbage neighbors under ambient ozone were less susceptible to oviposition by P. brassicae in the field, but not in the laboratory. At ambient ozone levels P. xylostella preferred to oviposit on cabbage plants exposed to undamaged conspecifics in both the field and laboratory. At enriched ozone levels in the field, P. xylostella showed no oviposition preference for cabbage exposed to damaged or undamaged cabbage neighbors. In multiple choice tests in the laboratory, P. xylostella preferred to oviposit on cabbage plants exposed to damaged broccoli plants at ambient ozone over plants exposed to undamaged broccoli plants at ambient ozone and damaged and undamaged broccoli plants at enriched ozone. We conclude that plant–plant interactions and ozone interplay when influencing the susceptibility of plants to oviposition by herbivores.

Volatiles of Solena amplexicaulis (Lam.) Gandhi Leaves Influencing Attraction of Two Generalist Insect Herbivores.

Epilachna vigintioctopunctata Fabr. (Coleoptera: Coccinellidae) and Aulacophora foveicollis Lucas (Coleoptera: Chrysomelidae) are important pests of Solena amplexicaulis (Lam.) Gandhi (Cucurbitaceae), commonly known as creeping cucumber. The profiles of volatile organic compounds from undamaged plants, plants after 48hr continuous feeding of adult females of either E. vigintioctopunctata or A. foveicollis, by adults of both species, and after mechanical damaging were identified and quantified by GC-MS and GC-FID analyses. Thirty two compounds were detected in volatiles of all treatments. In all plants, methyl jasmonate was the major compound. In Y-shaped glass tube olfactometer bioassays under laboratory conditions, both insect species showed a significant preference for complete volatile blends from insect damaged plants, compared to those of undamaged plants. Neither E. vigintioctopunctata nor A. foveicollis showed any preference for volatiles released by heterospecifically damaged plants vs. conspecifically damaged plants or plants attacked by both species. Epilachna vigintioctopunctata and A. foveicollis showed attraction to three different synthetic compounds, linalool oxide, nonanal, and E-2-nonenal in proportions present in volatiles of insect damaged plants. Both species were attracted by a synthetic blend of 1.64μg linalool oxide + 3.86μg nonanal + 2.23μg E-2-nonenal, dissolved in 20μl methylene chloride. This combination might be used as trapping tools in pest management strategies.

Herbivore Damage and Prior Egg Deposition on Host Plants Influence the Oviposition of the Generalist Moth Trichoplusia ni (Lepidoptera: Noctuidae).

Female insects have the difficult task of locating host plants that maximize the survival and success of their offspring. In this study, the oviposition preferences of the cabbage looper moth, Trichoplusia ni (Hübner), for soybean plants, Glycine max (L.), under various treatments-undamaged, mechanically damaged, damaged by T. ni or Spodoptera frugiperda (Smith) larvae or by Bemisia tabaci (Gennadius) adults, egg-free plants, and plants previously oviposited by conspecific or heterospecific females (S. frugiperda)-were investigated using two-choice tests. Additionally, the volatile compounds emitted by the plants under the different treatments were identified by gas chromatography-mass spectrometry. Our results showed that females showed no preferences for undamaged or mechanically damaged plants. However, they oviposited more often on undamaged plants than on those previously damaged by T. ni, S. frugiperda, or B. tabaci. In contrast, females preferred to oviposit on plants previously oviposited by conspecific and heterospecific females than on egg-free plants. Plants damaged by conspecific or heterospecific larvae emitted methyl salicylate, indole, and octyl butyrate, compounds not released by undamaged or mechanically damaged plants. Whitefly damage induced the release of higher quantities of Z(3)-hexenyl acetate, (R)-(+)-limonene, and (E)-β-ocimene compared to plants damaged by larvae and suppressed the emission of linalool. Egg deposition by conspecific and heterospecific moths induced the emission of (R)-(+)-limonene, octyl butyrate, and geranyl acetone but suppressed the release of linalool. This study showed that a generalist moth species can discriminate between plants of different quality, and suggests that females use volatile compounds as cues during this process.

Latitudinal Gradients in Induced and Constitutive Resistance against Herbivores

Plants are hypothesized to evolve increased defense against herbivores at lower latitudes, but an increasing number of studies report evidence that contradicts this hypothesis. Few studies have examined the evolution of constitutive and induced resistance along latitudinal gradients. When induction is not considered, underlying patterns of latitudinal clines in resistance can be obscured because plant resistance represents a combination of induced and constitutive resistance, which may show contrasting patterns with latitude. Here, we asked if there are latitudinal gradients in constitutive versus induced resistance by using genotypes of Oenothera biennis (Onagraceae) sampled along an 18° latitudinal gradient. We conducted two bioassay experiments to compare the resistance of plant genotypes against one generalist (Spodoptera exigua) and one specialist (Acanthoscelidius acephalus) herbivore. These insects were assayed on: i) undamaged control plants, ii) plants that had been induced with jasmonic acid, and iii) plants induced with herbivore damage. Additionally, we examined latitudinal gradients of constitutive and induced chemical resistance by measuring the concentrations of total phenolics, the concentration of oxidized phenolics, and the percentage of phenolics that were oxidized. Spodoptera exigua showed lower performance on plants from lower latitudes, whereas A. acephalus showed no latitudinal pattern. Constitutive total phenolics were greater in plants from lower latitudes, but induced plants showed higher total phenolics at higher latitudes. Oxidative activity was greatest at higher latitudes regardless of induction. Overall, both latitude and induction have an impact on different metrics of plant resistance to herbivory. Further studies should consider the effect of induction and herbivore specialization more explicitly, which may helpto resolve the controversy in latitudinal gradients in herbivory and defense.

Attraction of Telenomus podisi to volatiles induced by Euschistus heros in three different plant species

Specialized natural enemies that forage for polyphagous hosts need to locate hosts on different plants. Telenomus podisi (Hymenoptera: Platygastridae) is a stink bug egg parasitoid with a preference for Euschistus heros (Hemiptera, Pentatomidae), a polyphagous species. The aim of this study was to evaluate the induction of defences in three E. heros host plants: maize (Zea mays), sunflower (Helianthus annuus) and pigeon pea (Cajanus cajan). We hypothesized that E. heros damage to these three plants enhances the attraction of the parasitoid T. podisi as has been observed in other systems. Using Y-tube olfactometer bioassays, we tested parasitoid responses to combinations of the following odour sources: clean air, undamaged plants and plants damaged by stink bug feeding. Volatiles were collected by means of dynamic headspace collection and analysed by gas chromatography coupled to mass spectrometry. T. podisi did not distinguish odours from undamaged plants against air for any of the three plant species. For maize, the parasitoid preferred the odour from herbivore-damaged plants over both clean air and undamaged plants. For sunflower, the parasitoid only preferred the odour of herbivore-damaged plants over the odour of undamaged plants. For pigeon pea, no preferences were observed. Quantitative differences in the volatile profile of damaged and undamaged plants were observed in each plant species. We conclude that sunflower and maize plants, when damaged by E. heros, release volatiles that attract the parasitoid T. podisi; the parasitoid appears to use a different blend composition to distinguish herbivore-damaged plants of each species.

Stage-Related Defense Response Induction in Tomato Plants by Nesidiocoris tenuis

The beneficial effects of direct predation by zoophytophagous biological control agents (BCAs), such as the mirid bug Nesidiocoris tenuis, are well-known. However, the benefits of zoophytophagous BCAs’ relation with host plants, via induction of plant defensive responses, have not been investigated until recently. To date, only the females of certain zoophytophagous BCAs have been demonstrated to induce defensive plant responses in tomato plants. The aim of this work was to determine whether nymphs, adult females, and adult males of N. tenuis are able to induce defense responses in tomato plants. Compared to undamaged tomato plants (i.e., not exposed to the mirid), plants on which young or mature nymphs, or adult males or females of N. tenuis fed and developed were less attractive to the whitefly Bemisia tabaci, but were more attractive to the parasitoid Encarsia formosa. Female-exposed plants were more repellent to B. tabaci and more attractive to E. formosa than were male-exposed plants. When comparing young- and mature-nymph-exposed plants, the same level of repellence was obtained for B. tabaci, but mature-nymph-exposed plants were more attractive to E. formosa. The repellent effect is attributed to the signaling pathway of abscisic acid, which is upregulated in N. tenuis-exposed plants, whereas the parasitoid attraction was attributed to the activation of the jasmonic acid signaling pathway. Our results demonstrate that all motile stages of N. tenuis can trigger defensive responses in tomato plants, although these responses may be slightly different depending on the stage considered.

An Indirect Defence Trait Mediated through Egg-Induced Maize Volatiles from Neighbouring Plants.

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.

Effects of single and dual species herbivory on the behavioral responses of three thrips species to cotton seedlings.

This study investigated the olfactory responses of 3 thrips species [Frankliniella schultzei Trybom, F. occidentalis Pergrande and Thrips tabaci Lindeman (Thysanoptera: Thripidae)] to cotton seedlings [Gossypium hirsutum L. (Malvales: Malvaceae)] simultaneously damaged by different combinations of herbivores. Cotton seedlings were damaged by foliar feeding Tetranychus urticae Koch (Trombidiforms: Tetranychidae), Helicoverpa armigera Hübner (Lepidoptera: Noctuidae), Aphis gossypii Glover (Hemiptera: Aphididae) or root feeding Tenebrio molitor L. (Coleoptera: Tenebrionidae). Thrips responses to plants simultaneously damaged by 2 species of herbivore were additive and equivalent to the sum of the responses of thrips to plants damaged by single herbivore species feeding alone. For example, F. occidentalis was attracted to T. urticae damaged plants but more attracted to undamaged plants than to plants damaged by H. armigera. Plants simultaneously damaged by low densities of T. urticae and H. armigera repelled F. occidentalis but as T. urticae density increased relative to H. armigera density, F. occidentalis attraction to coinfested plants increased proportionally. Thrips tabaci did not discriminate between undamaged plants and plants damaged by H. armigera but were attracted to plants damaged by T. urticae alone or simultaneously damaged by T. urticae and H. armigera. Olfactometer assays showed that simultaneous feeding by 2 herbivores on a plant can affect predator-prey interactions. Attraction of F. occidentalis to plants damaged by its T. urticae prey was reduced when the plant was simultaneously damaged by H. armigera, T. molitor, or A. gossypii and F. schultzei was more attracted to plants simultaneously damaged by T. urticae and H. armigera than to plants damaged by T. urticae alone. We conclude that plant responses to feeding by 1 species of herbivore are affected by responses to feeding by other herbivores. These plant-mediated interactions between herbivore complexes affect the behavioral responses of thrips which vary between species and are highly context dependent.

Effects of florivory on plant-pollinator interactions: Implications for male and female components of plant reproduction.

Florivory could have direct negative effects on plant fitness due to consumption of floral organs, and indirect effects mediated through changes in traits important to pollination. These effects likely vary with plant sexual system, depending on sex- or morph-specific patterns of damage. We investigated the direct and indirect effects of simulated florivory on male and female components of reproduction in the native, distylous vine Gelsemium sempervirens. We crossed floral damage and supplemental pollination treatments in a common garden array and tracked pollinator behavioral responses. We also estimated male function using fluorescent dye as an analog for pollen transfer, and measured both fruit and seed production. The effects of floral damage varied by floral morph, the genus of floral visitor, and the component of reproduction measured. Damage reduced the number of pollinator visits to pin but not thrum plants, and increased the time some pollinators spent per flower in thrum but not pin plants. Flowers of damaged plants transferred more dye particles to recipient plants compared to undamaged plants, but only later in the season when the majority of dye transfer occurred. Damage had no effect on female reproduction. These results suggest that florivory can have positive indirect effects on estimated male plant reproduction through changes in different pollinators' behavior at flowers, but the effects of floral damage vary with male vs. female function. These results underscore the importance of other species' interactions at flowers in driving pollinator behavior and pollen transfer dynamics.

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.

Elevated Ozone Modulates Herbivore-Induced Volatile Emissions of Brassica nigra and Alters a Tritrophic Interaction.

Plants damaged by herbivores emit volatile organic compounds (VOCs) that are used by parasitoids for host location. In nature, however, plants are exposed to multiple abiotic and biotic stresses of varying intensities, which may affect tritrophic interactions. Here, we studied the effects of ozone exposure and feeding by Pieris brassicae larvae on the VOCs emitted by Brassica nigra and the effects on oriented flight of the parasitoid Cotesia glomerata. We also investigated the oriented flight of C. glomerata in a wind-tunnel with elevated ozone levels. Herbivore-feeding induced the emission of several VOCs, while ozone alone had no significant effect. However, exposure to 120ppb ozone, followed by 24hr of herbivore-feeding, induced higher emissions of all VOCs as compared to herbivore-feeding alone. In accordance, herbivore-damaged plants elicited more oriented flights than undamaged plants, whereas plants exposed to 120ppb ozone and 24hr of herbivore-feeding elicited more oriented flights than plants subjected to herbivore-feeding alone. Ozone enrichment of the wind-tunnel air appeared to negatively affect orientation of parasitoids at 70ppb, but not at 120ppb. These results suggest that the combination of ozone and P. brassicae-feeding modulates VOC emissions, which significantly influence foraging efficiency of C. glomerata.

Feeding Experience Affects the Behavioral Response of Polyphagous Gypsy Moth Caterpillars to Herbivore-induced Poplar Volatiles.

Plant volatiles influence host selection of herbivorous insects. Since volatiles often vary in space and time, herbivores (especially polyphagous ones) may be able to use these compounds as cues to track variation in host plant quality based on their innate abilities and previous experience. We investigated the behavioral response of naïve (fed on artificial diet) and experienced (fed on poplar) gypsy moth (Lymantria dispar) caterpillars, a polyphagous species, towards constitutive and herbivore-induced black poplar (Populus nigra) volatiles at different stages of herbivore attack. In Y-tube olfactometer assays, both naïve and experienced caterpillars were attracted to constitutive volatiles and volatiles released after short-term herbivory (up to 6 hr). Naïve caterpillars also were attracted to volatiles released after longer-term herbivory (24–30 hr), but experienced caterpillars preferred the odor of undamaged foliage. A multivariate statistical analysis comparing the volatile emission of undamaged plants vs. plants after short and longer-term herbivory, suggested various compounds as being responsible for distinguishing between the odors of these plants. Ten compounds were selected for individual testing of caterpillar behavioral responses in a four-arm olfactometer. Naïve caterpillars spent more time in arms containing (Z)-3-hexenol and (Z)-3-hexenyl acetate than in solvent permeated arms, while avoiding benzyl cyanide and salicyl aldehyde. Experienced caterpillars avoided benzyl cyanide and preferred (Z)-3-hexenyl acetate and the homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) over solvent. Only responses to DMNT were significantly different when comparing experienced and naïve caterpillars. The results show that gypsy moth caterpillars display an innate behavioral response towards constitutive and herbivore-induced plant volatiles, but also that larval behavior is plastic and can be modulated by previous feeding experience.Electronic supplementary materialThe online version of this article (doi:10.1007/s10886-016-0698-7) contains supplementary material, which is available to authorized users.

Susceptibility of Cultivated Plants to Sumatran Elephant (Elephas maximus sumatranus) in The Human Elephants Conflict Area in Aceh Province

Study on human-elephant conflict was conducted in Aceh Province in August 2013−April 2014 to assess susceptibility of farms by crop raiding elephants. The locations were determined by selected areas impacted by elephant conflict including Cot Girek, Mane, Meureudu, Sampoiniet, and Pantai Ceureumen. 150 respondents was interviewed to assess the variety of the commodity plant species cultivated by local community within study areas, species of damaged commodity plants, species of undamaged commodity plants, and the planting system. There were 29 species considered as commodity plants cultivated by farmers. Moreover, 5 commodity plants were considered as high risk plants damaged by elephant including areca, banana, oil palm, paddy, and rubber. On the other hand, species considered as low risk or undamaged consist of cacao, coffee, chili, candlenut, and patchioli. Those low risk or undemaged commodity plants species have a potential to be promoted as elephant-friendly crop commodities in area adjacent to elephant habitat based on the analysis and the categorization of susceptibility of cultivated plants against crop raiding elephant. One of the problems of human-elephant conflict is crop raiding of village farms. It is assumed that elephants might destroy a particular species therefore information on the species could assist farmers in selecting appropriate crop to be planted. There is a risk that current polyculture and monoculture planting system used by farmers will not prevent farms from crop raiding elephants.

Cross-modal effect of natal habitat experience increases receptivity to non-natal habitat cues in generalist parasitic wasps

Exposure to natal habitats induces preferences in individual organism for foraging cues that originate from their developmental habitat. However, the natal experience of habitat generalists may play a different role in their habitat selection, since they use a broad range of foraging cues in a non-natal habitat. In this study, the effects of natal habitat experiences on the responses of females of the generalist parasitoid Aphidius gifuensis Ashmead to cues from natal or non-natal habitats were investigated. The landing rates on plant-host complexes (PHCs) and undamaged plants (UDPs) of natal (wheat) and non-natal (broad bean) plant systems in a wind tunnel increased after encounters with host aphids on the wheat-PHCs when the wasps had previously been exposed to wheat-UDPs. However, without the previous exposure to wheat-UDPs, the landing rate on the broad bean-PHCs was not increased. Similar increase found in responses to green paper disc dummies suggested involvement of visual stimuli in the reinforcement. In olfactometer tests, preferences for broad bean-UDPs over broad bean-PHCs were found but wasps with previous exposure to wheat-UDPs and host encounter on wheat-PHCs did not show the preference. These results suggest that the early exposure to natal habitat plants modified later learning and response to visual cues from potential non-natal habitat plants, and olfactory learning might be involved in the modification. This cross-modal effect of early natal experience would compensate for the loss of host-searching efficiency due to uncertain information of potential non-natal habitats.

Low tolerance to simulated herbivory in Hawaiian seedlings despite induced changes in photosynthesis and biomass allocation.

Seedling herbivory is an important factor underlying plant community diversity and structure. While considerable research has characterized seedling defence in terms of resistance, very little is known about seedling tolerance of herbivory. Moreover, few studies have attempted to identify mechanisms of tolerance across a range of plant species. Seedling tolerance of simulated herbivory was tested in a diverse pool of ten Hawaiian plant species, including several lobeliad species (family Campanulaceae), a grass, a herb and common woody trees and shrubs. Tolerance was measured as the relative survival and growth of damaged plants receiving 50% defoliation with simultaneous jasmonic acid application compared with undamaged control plants, assessed 1·5 and 5weeks after damage. Putative mechanisms of tolerance were measured, including photosynthetic parameters, light use efficiency, and biomass allocation reflecting growth priorities, and analysed using species-level regression analyses on tolerance indices. No species fully tolerated 50% defoliation at either harvest date, and simulated herbivory significantly reduced shoot as well as root biomass. Lobeliad species had particularly low tolerance. Species varied considerably in size, biomass allocation parameters and their constitutive (pre-damage) and induced (post-damage) photosynthetic parameters. However, only constitutive levels of non-photochemical quenching were significantly related to tolerance, indicating that species with more efficient light use (and less heat dissipation) are better at tolerating damage than species with high levels of heat dissipation. Native Hawaiian plants expressed low tolerance to a conservative level of simulated herbivory. Root growth decreased in response to damage, but this was not associated with greater tolerance, suggesting this response may be due to allocation constraints following defoliation and not due to adaptive plasticity. Conservation of native island plants threatened by invasive herbivores should prioritize protection for seedlings for improved regeneration and the persistence of native plants in disturbed habitats.

Tolerance of Swallowworts (Vincetoxicumspp.) to Multiple Years of Artificial Defoliation and Clipping

AbstractThe European vines pale swallowwort and black swallowwort are invading various habitats in northeastern North America. It is unclear how these plants might respond to potential biological control agents, as they experience little herbivore damage in North America, or longer durations of mowing given the reported lack of efficacy of mechanical control. We evaluated the effect of six seasons of artificial defoliation (50 or 100% defoliation once or twice per season) and clipping (once, twice, or four times at 8 cm above the soil level) on the survival, growth, and reproduction of mature plants of the two species grown in a common garden field experiment. No plants died from damage after 6 yr. Black swallowwort produced more aboveground biomass, whereas pale swallowwort produced more root biomass and root crown buds, compared with its congener species. For most damage treatments, root biomass and the number of crown buds and stems increased over time, whereas aboveground biomass and viable seeds per plant generally did not change. Substantial overlap in plant size and seed production occurred among damage treatments and species. The most severe defoliation treatment did not substantially limit growth and reproduction compared with undamaged plants. While two clippings per season sometimes prevented seed production, four clippings per season was the only type of damage that consistently prevented plant growth and eliminated seed production. Pale and black swallowwort display a high tolerance to aboveground tissue loss in high-light environments without plant competition. The annual increase in plant size calls into question the potential efficacy of a defoliating insect against field populations of swallowworts, and it seems likely the only benefits of a long-term mowing regime will be to eliminate seed production.

Characterization of Feeding Injuries Caused by Ceresa nigripectus Remes Lenicov (Hemiptera: Membracidae) on Alfalfa Stems.

Piercing-sucking insects cause mechanical and physiological injury to plants. Ceresa nigripectus Remes Lenicov is a pest of alfalfa in subtropical regions of South America and a carrier of the ArAWB phytoplasma. The aim of this study was to determine the feeding habits of this treehopper and to describe the effects of the feeding injuries on stem vascular tissues in alfalfa. Adults and nymphs of C. nigripectus inserted their stylets repeatedly girdling the stem. One week after feeding, alfalfa stems exhibited numerous feeding canals with salivary deposits, most of which reached the phloem. Two weeks after feeding, cortex and phloem cells next to the salivary sheath collapsed, mature tracheal elements became sparse and appeared with an increased cross-section area, and phenolic compounds increased in cells and cell walls compared to undamaged plants. Three weeks after feeding, an annular callus, formed by abnormal cell division and hypertrophy of preexisting cortex and vascular cambium cells, appeared immediately above the stem girdle. Parenchyma cells from the outer layers of the callus differentiated to form secondary anomalous amphicribal bundles in the wound. The aerial parts above the stem girdle eventually withered and died.