森林生态系统中植食性昆虫与寄主的互作机制、假说与证据

围绕"多样性稳定性"假说、"联合抗性假说"、"生长势假说"、"胁迫假说"、以及下调、上调和推拉等机制与假说提出的背景与实验验证的证据,力图辨析其概念以及它们之间的相互关系。作者认为,多样性-稳定性机制关注森林生态系统的功能,是基于群落甚至景观层次。多样性条件下的联合抗性机制和联合易感性应属于稳定性中的抵抗力范畴。联合抗性机制的主要基础是基于资源集中假说和天敌假说,这些观点在种群层次上更易理解;上调力和下调力机制是以食物网底部的资源与顶端的天敌来探讨这种互作关系。因此,资源集中与上调力有着对应关系,而天敌假说只是下调力机制中的一个层面而已。植物生长势假说和植物胁迫假说力图从植物个体或种的群体的生长状态出发解析植食性动物的对寄主的选择趋势。上述有关植食性昆虫与寄主互作的机制、假说与证据是基于不同的层面提出的,因而在解析研究目标时,由于基本面的差异有可能会得出不同的结论。以近年来的研究进展和研究成果为依据有针对性地阐述这些理论对森林有害生物生态调控技术的指导作用,其中,联合抗性和联合易感性理论对指导森林有害生物生态控制具有更直接的指导作用。进一步提出了相应的亟待解决的科学问题。     

Non-host volatiles mediate associational resistance to the pine processionary moth

An increasing body of evidence indicates that the association between different plant species may lead to a reduction in insect herbivory, i.e. associational resistance. This might be due to a top–down regulation of herbivores by increased numbers of natural enemies or to a disruptive bottom–up influence of lower host plant accessibility. In particular, the richer plant communities release more diverse plant odours that may disturb olfactory-guided host choice and mating behaviour of insect herbivores, i.e. the “semiochemical diversity hypothesis”. However, this hypothesis has been rarely tested experimentally in natural habitats, notably forest ecosystems. We tested the effects of non-host volatiles (NHV) on mate and host location by the pine processionary moth (PPM) at the scale of individual pine trees with branches of non-host tree (birch) at their base. Pheromone trap catches and the numbers of larval nests were both reduced by non-host presence under treated pine trees, confirming an associational resistance mediated by NHV. In both males and females, the antenna could detect several birch volatiles, including methyl salicylate (MeSa). MeSa inhibited the attraction of the PPM male to pheromone traps, as did bark and leaf chips from birch trees. Our test of three doses of MeSa at the habitat scale (50 m forest edges) showed that the reduction in the numbers of male PPM captured in traps and in larval nests was MeSa dose-dependent. These results show that odours released by deciduous non-host trees can reduce herbivory by a forest defoliator in conifers, providing support to the “semiochemical diversity hypothesis” as a mechanism of associational resistance.     

Host plant predictability and the feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores

Summary Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.     

Host plant scents attract rolled-leaf beetles to Neotropical gingers in a Central American tropical rain forest

Leaf volatile chemicals are known to reduce herbivory rates by repelling or intoxicating insect herbivores and by attracting the predators and parasitoids of herbivores. However, leaf volatiles may also be used by insect herbivores as cues to locate their host plants. Leaf volatiles are suggested to be important host search cues for herbivores in structurally complex and diverse habitats, such as tropical rain forests. A group of insect herbivores, the rolled-leaf beetles (Coleoptera: Chrysomelidae: Hispinae), have maintained a highly specialized interaction with Neotropical gingers (Zingiberales) for ca. 60 million years. In this study, we explored chemical attraction to host plants under controlled laboratory conditions, using four sympatric rolled-leaf beetle species, Cephaloleia dorsalis Baly, Cephaloleia erichsonii Baly, Cephaloleia fenestrata Weise, and Cephaloleia placida Baly. For each beetle species, we investigated (i) whether it was repelled or attracted by leaf scents produced by four host and four non-host plant species, including Neotropical gingers in the families Marantaceae, Costaceae, and Zingiberaceae; and (ii) its ability to use scents to detect its host plant. We found that rolled-leaf beetles can detect and are attracted by leaf volatiles from both host and non-host gingers. Additionally, when beetles were simultaneously exposed to leaf volatiles from host and non-host plants, three rolled-leaf beetle species were significantly more attracted by volatiles from their host plants than from non-hosts. Only one of the beetle species was not able to discriminate between host and non-host scents.     

A quantitative evaluation of major plant defense hypotheses, nature versus nurture, and chemistry versus ants

Variation in plant secondary metabolite content can arise due to environmental and genetic variables. Because these metabolites are important in modifying a plant’s interaction with the environment, many studies have examined patterns of variation in plant secondary metabolites. Investigations of chemical defenses are often linked to questions about the efficacies of plant defenses and hypotheses on their evolution in different plant guilds. We performed a series of meta-analyses to examine the importance of environmental and genetic sources of variation in secondary metabolites as well as the antiherbivore properties of different classes of defense. We found both environmental and genetic variation affect secondary metabolite production, supporting continued study of the carbon-nutrient balance and growth-differentiation balance hypotheses. Defenses in woody plants are more affected by genetic variation, and herbaceous plant defenses are more influenced by environmental variation. Plant defenses in agricultural and natural systems show similar responses to manipulations, as do plants in laboratory, greenhouse, or field studies. What does such variation mean to herbivores? A comparison of biotic, physical, and chemical defenses revealed the most effective defensive strategy for a plant is biotic mutualisms with ants. Fast-growing plants are most often defended with qualitative defenses and slow-growing plants with quantitative defenses, as the plant apparency and resource availability hypotheses predict. However, we found the resource availability hypothesis provides the best explanation for the evolution of plant defenses, but the fact that there is considerable genetic and environmental variation in defenses indicates herbivores can affect plant chemistry in ecological and evolutionary time.     

Colorado potato beetle manipulates plant defenses in local and systemic leaves

Herbivore microbial associates can affect diverse interactions between plants and insect herbivores. Some insect symbionts enable herbivores to expand host plant range or to facilitate host plant use by modifying plant physiology. However, little attention has been paid to the role of herbivore-associated microbes in manipulating plant defenses. We have recently shown that Colorado potato beetle secrete the symbiotic bacteria to suppress plant defenses. The bacteria in oral secretions from the beetle hijack defense signaling pathways of host plants and the suppression of induced plant defenses benefits the beetle’s performance. While the defense suppression by the beetle-associated bacteria has been investigated in local damaged leaves, little is known about the effects of the symbiotic bacteria on the manipulation of plant defenses in systemic undamaged leaves. Here, we demonstrate that the symbiotic bacteria suppress plant defenses in both local and systemic tissues when plants are attacked by antibiotic-untreated larvae.     

Going undercover: increasing canopy cover around a host tree drives associational resistance to an insect pest

Neighbouring heterospecific plants are often observed to reduce the probability of herbivore attack on a given focal plant. While this pattern of associational resistance is frequently reported, experimental evidence for underlying mechanisms is rare particularly for potential plant species diversity effects on focal host plants and their physical environment. Here, we used an established forest diversity experiment to determine whether tree diversity effects on an important insect pest are driven by concomitant changes in host tree growth or the light environment. We examined the effects of tree species richness, canopy cover and tree growth on the probability of occurrence, the abundance, and volume of galls caused by the pineapple gall adelgid Adelges abietis on Norway spruce. Although tree diversity had no effect on gall abundance, we observed that both the probability of gall presence and gall volume (an indicator of maternal fecundity) decreased with tree species richness and canopy cover around host spruce trees. Structural equation models revealed that effects of tree species richness on gall presence and volume were mediated by concurrent increases in canopy cover rather than changes in tree growth or host tree density. As canopy cover did not influence tree or shoot growth, patterns of associational resistance appear to be driven by improved host tree quality or more favourable microclimatic conditions in monocultures compared to mixed‐stands. Our study therefore demonstrates that changes in forest structure may be critical to understanding the responses of herbivores to plant diversity and may underpin associational effects in forest ecosystems.     

Non-native plants rarely provide suitable habitat for native gall-inducing species

For insect herbivores, a critical niche requirement—possibly the critical niche requirement—is the presence of suitable host plants. Current research suggests that non-native plants are not as suitable as native plants for native herbivores, resulting in decreases in insect abundance and richness on non-native plants. Like herbivores, gall-forming insects engage in complex, species-specific interactions with host plants. Galls are plant tissue tumors (including bulbous or spindle-shaped protrusions on leaves, stems and other plant organs) that are induced by insects through physical or chemical damage (prompting plants to grow a protective tissue shell around the insect eggs and larvae). As such, we hypothesized that gall-inducing insect species richness would be higher on native than non-native plants. We also predicted higher gall-inducing insect species richness on woody than herbaceous plants. We used an extensive literature review in which we compiled gall host plant species by genus, and we assigned native or non-native (or mixed) status to each genus. We found that native plants host far more gall-inducing insect species than non-native plants; woody plants host more gall-inducing species than herbaceous plants; and native woody plants host the most gall-inducing species of all. Gall-inducing species generally are a very cryptic group, even for experts, and hence do not elicit the conservation efforts of more charismatic insects such as plant pollinators. Our results suggest that non-native plants, particularly non-native woody species, diminish suitable habitat for gall-inducing species in parallel with similar results found for other herbivores, such as Lepidopterans. Hence, the landscape-level replacement of native with non-native species, particularly woody ones, degrades taxonomically diverse gall-inducing species (and their inquilines and parasitoids), removing multiple layers of diversity from forest ecosystems.

     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

Aggregative oviposition behaviour of Helicoverpa spp. (Lepidoptera: Noctuidae) in contaminated chickpea crops

Abstract Aggregative egg distribution by Helicoverpa spp. within experimental field plots of chickpea containing various contaminant plant species was documented at different stages of plant growth. More eggs were laid on contaminant faba and wheat plants than on the surrounding chickpea. The level of egg aggregation on faba and wheat plants increased with general crop growth, becoming evident before a significant disparity in growth rates between these species and the surrounding chickpea became apparent. In chickpea plots contaminated with canola, aggregative oviposition activity on the contaminant plants was observed in the early seedling stage whilst a clear height differential between the two species was evident. Total Helicoverpa egg density on contaminant plants was not correlated with height above chickpea for any of the contaminant species. It is proposed that the aggregative oviposition behaviour of Helicoverpa spp. in contaminated chickpea results from differential apparency of the contaminant plants to ovipositing moths. Differential apparency may result from several factors including growth rate and morphological differences between chickpea and the contaminants. Physiological changes in chickpea, including growth stage-related changes in acid production by the foliage, may also contribute to the greater apparency of contaminant plants in chickpea plots. The implications of differential host plant apparency for insect pest management are discussed.     

Land‐use legacies and present fire regimes interact to mediate herbivory by altering the neighboring plant community

Past and present human activities, such as historic agriculture and fire suppression, are widespread and can create depauperate plant communities. Although many studies show that herbivory on focal plants depends on the density of herbivores or the composition of the surrounding plant community, it is unclear whether anthropogenic changes to plant communities alter herbivory. We tested the hypothesis that human activities that alter the plant community lead to subsequent changes in herbivory. At 20 sites distributed across 80 300 hectares, we conducted a field experiment that manipulated insect herbivore access (full exclosures and pseudo‐exclosures) to four focal plant species in longleaf pine woodlands with different land‐use histories (post‐agricultural sites or non‐agricultural sites) and degrees of fire frequency (frequent and infrequent). Plant cover, particularly herbaceous cover, was lower in post‐agricultural and fire suppressed woodlands. Density of the dominant insect herbivore at our site (grasshoppers) was positively related to plant cover. Herbivore access reduced biomass of the palatable forb Solidago odora in frequently burned post‐agricultural sites and in infrequently burned non‐agricultural woodlands and increased mortality of another forb (Pityopsis graminifolia), but did not affect two other less palatable species (Schizachyrium scoparium and Tephrosia virginiana). Herbivory on S. odora exhibited a hump‐shaped response to plant cover, with low herbivory at low and high levels of plant cover. Herbivore density had a weak negative effect on herbivory. These findings suggest that changes in plant cover related to past and present human activities can modify damage rates on focal S. odora plants by altering grasshopper foraging behavior rather than by altering local grasshopper density. The resulting changes in herbivory may have the potential to limit natural recovery or restoration efforts by reducing the establishment or performance of palatable plant species.     

Non-host plant extracts reduce oviposition of Plutella xylostella (Lepidoptera: Plutellidae) and enhance parasitism by its parasitoid Cotesia plutellae (Hymenoptera: Braconidae)

Botanical preparations, usually from non-host plants, can be used to manipulate the behaviour of insect pests and their natural enemies. In this study, the effects of extracts of Chrysanthemum morifolium, a non-host plant of the diamondback moth, Plutella xylostella (Linnaeus), on the olfactory and oviposition responses of this phytophagous insect and on levels of parasitism by its specialist parasitoid Cotesia plutellae (Kurdjumov) were examined, using Chinese cabbage Brassica campestris L. ssp. pekinensis as the test host plant. Olfactometer tests showed that volatiles of chrysanthemum extract-treated host plants were less attractive to P. xylostella females than those from untreated host plants; and in contrast, volatiles of the chrysanthemum extract-treated host plants were more attractive to females of its parasitoid C. plutellae than those from untreated host plants. Oviposition preference tests showed that P. xylostella females laid only a small proportion of their eggs on chrysanthemum extract-treated host plants, while ovipositing parasitoid females parasitized a much higher proportion of host larvae feeding on the treated host plants than on untreated host plants. These results suggest that certain non-host plant compounds, when applied onto a host plant, may render the plant less attractive to a phytophagous insect but more attractive to its parasitoids. Application of such non-host plant compounds can be explored to develop push-pull systems to reduce oviposition by a pest insect and at the same time enhance parasitism by its parasitoids in crops.     

APHID DISTRIBUTION AND THE EVOLUTION OF GOLDENROD RESISTANCE

Although there is considerable evidence indicating that herbivory is detrimental to plant fitness, some recent studies of the evolution of plant resistance have concluded that insects do not impose selection on their host plants. A previously untested assumption that underlies most studies of the evolution of plant resistance is that insect distribution patterns are controlled directly by the effects of plant genotype on insect preference and performance. The experiments described here explicitly tested this assumption using the specialist herbivore Uroleucon tissoti (Homoptera: Aphididae) and its host plant Solidago altissima (Asteraceae). Measures of aphid preference and performance were used to predict aphid distribution patterns, and then the predicted distribution patterns were compared with the natural distribution pattern. Although goldenrod genotype had a strong effect on aphid distribution, aphid distribution was not controlled directly by the effect of goldenrod genotype on aphid preference and performance. Instead, a second experiment demonstrated that aphid and spittlebug (Philaenus spumarius and Lepyronia quadrangularis Homoptera: Cercopidae) distribution is controlled largely by genetic variation for resistance to a suite of “branch-causing” herbivores. These herbivores induce branching and aphids and spittlebugs are more abundant on branched plants than unbranched plants. These results indicate that any natural selection imposed by aphids and spittlebugs on goldenrod will depend on the presence or absence of branch-causing herbivores. Thus, selection for plant resistance may depend as much on the assemblage of insect species present as on the identity of each individual species.     

Habitats as Complex Odour Environments: How Does Plant Diversity Affect Herbivore and Parasitoid Orientation?

Plant diversity is known to affect success of host location by pest insects, but its effect on olfactory orientation of non-pest insect species has hardly been addressed. First, we tested in laboratory experiments the hypothesis that non-host plants, which increase odour complexity in habitats, affect the host location ability of herbivores and parasitoids. Furthermore, we recorded field data of plant diversity in addition to herbivore and parasitoid abundance at 77 grassland sites in three different regions in Germany in order to elucidate whether our laboratory results reflect the field situation. As a model system we used the herb Plantago lanceolata, the herbivorous weevil Mecinus pascuorum, and its larval parasitoid Mesopolobus incultus. The laboratory bioassays revealed that both the herbivorous weevil and its larval parasitoid can locate their host plant and host via olfactory cues even in the presence of non-host odour. In a newly established two-circle olfactometer, the weeviĺs capability to detect host plant odour was not affected by odours from non-host plants. However, addition of non-host plant odours to host plant odour enhanced the weeviĺs foraging activity. The parasitoid was attracted by a combination of host plant and host volatiles in both the absence and presence of non-host plant volatiles in a Y-tube olfactometer. In dual choice tests the parasitoid preferred the blend of host plant and host volatiles over its combination with non-host plant volatiles. In the field, no indication was found that high plant diversity disturbs host (plant) location by the weevil and its parasitoid. In contrast, plant diversity was positively correlated with weevil abundance, whereas parasitoid abundance was independent of plant diversity. Therefore, we conclude that weevils and parasitoids showed the sensory capacity to successfully cope with complex vegetation odours when searching for hosts.     

Strong but opposing effects of associational resistance and susceptibility on defense phenotype in an African savanna plant

The susceptibility of plants to herbivores can be strongly influenced by the identity, morphology and palatability of neighboring plants. While the defensive traits of neighbors often determine the mechanism and strength of associational resistance and susceptibility, the effect of neighbors on plant defense phenotype remains poorly understood. We used field surveys and a prickle‐removal experiment in a semi‐arid Kenyan savanna to evaluate the efficacy of physical defenses against large mammalian herbivores in a common understory plant, Solanum campylacanthum. We then quantified the respective effects of spinescent Acacia trees and short‐statured grasses on browsing damage and prickle density in S. campylacanthum. We paired measurements of prickle density beneath and outside tree canopies with long‐term herbivore‐exclusion experiments to evaluate whether associational resistance reduced defense investment by decreasing browsing damage. Likewise, we compared defense phenotype within and outside pre‐existing and experimentally created clearings to determine whether grass neighbors increased defense investment via associational susceptibility. Removing prickles increased the frequency of browsing by ~25%, and surveys of herbivory damage on defended leaves suggested that herbivores tended to avoid prickles. As predicted, associational resistance and susceptibility had opposing effects on plant phenotype: individuals growing beneath Acacia canopies (or, analogously, within large‐herbivore exclosures) had a significantly lower proportion of their leaves browsed and produced ~ 70–80% fewer prickles than those outside refuges, whereas plants in grass‐dominated clearings were more heavily browsed and produced nearly twice as many prickles as plants outside clearings. Our results demonstrate that associational resistance and susceptibility have strong, but opposing, effects on plant defense phenotype, and that variable herbivore damage is a major source of intraspecific variation in defense phenotype in this system.     

Size does matter: variation in herbivory between and within plants and the plant vigor hypothesis

The plant stress and plant vigor hypotheses (PVH) are two of the most widely recognized hypothesis invoked to explain differential distribution of insect herbivores among their host plants. In both cases, the emphasis is on bottom–up processes (i.e. host-plant quality), but a recent meta-analytical review of the literature has shown that the plant stress hypothesis might have limited support among insect herbivores. In this study, we conducted a meta-analysis of the effects of plant vigor on insect herbivore abundance and survivorship by reviewing 71 published articles that explicitly tested the PVH and enabled 161 independent comparisons. Z-transform was used as the metric to standardize the results of all independent comparisons. Our quantitative results have shown that Hymenoptera (sawflies) was the most abundant group in the reviewed studies, representing 28.1% of the independent comparisons, followed by Diptera (25.1%) and Homoptera (22.6%). Amongst all the guilds studied, gall-formers were the most representative group (68.0%), whereas leaf-miners and stem-borers were underrepresented (less than 4.0% of the available comparisons). Insect herbivores were significantly more abundant on more vigorous plants (E++=0.6432, CI=0.7558–0.7280), but no significant effect was detected on herbivore survivorship. When herbivores were categorized into feeding guilds, effects of plant vigor on herbivore abundance were stronger for sap-suckers, leaf-miners and gall-formers. Our results have shown a strong herbivore preference for more vigorous plants, although our results do not support a preference–performance linkage.     

Influence of surrounding vegetation on insect herbivory: A matter of spatial scale and herbivore specialisation

The diversity of surrounding vegetation is thought to modify the interactions between a focal plant and its herbivores, disrupting (associational resistance) or enhancing (associational susceptibility) host plant location and colonisation. We compared the effects of host plant concentration on herbivory by generalist and specialist insects feeding on oak seedlings by increasing local concentration of seedlings. We also assessed the effects of the composition and structure of surrounding vegetation, both at stand and local levels. The damage caused by generalist leaf-feeding insects depended on the structure of plant communities at stand level, and increased with tree cover. By contrast, infestation by specialist leaf miners was affected by local understorey vegetation surrounding oak seedlings, and decreased with increasing shrub cover and stratification diversity. Leaf mine abundance was higher at higher oak seedling density, giving support to the host concentration hypothesis. However, the abundance of these specialist herbivores was also negatively correlated with damage caused by the generalist external leaf-feeders, suggesting competitive interactions.     

The deleterious effects of salinity stress on leafminers and their freshwater host

Abstract.  1. Salinity is an important cause of abiotic stress in wetland communities yet little is known about its consequences for freshwater plants and their insect herbivores. The goal of this study was to test the effect of salinity on a leafmining insect, Cerodontha iridiphora , and its herbaceous host plant, Iris hexagona .
2. Leafminer performance was evaluated on irises grown in control and saline treatments, and the effects of salinity and herbivory on leaf quality and mortality was measured.
3. Leafminer density and size were significantly lower on irises grown in saline water compared with freshwater.
4. Both salinity and herbivory accelerated leaf senescence and mortality, and their combined effects increased tissue loss by an order of magnitude compared with controls.
5. Leafminers acted as nutrient sinks. The undamaged regions of mined leaves contained 40% less nitrogen than unmined leaves, providing a mechanism for the premature leaf mortality.
6. Salinity was detrimental to the performance and survival of both the iris leafminer and its host plant. We propose that glycophytic host plants and their insect herbivores will suffer more than halophytic communities from environmental salinity because they lack the adaptive mechanisms to tolerate this potent physiological stress.