Insect herbivory and plant adaptation in an early successional community*

To address the role of insect herbivores in adaptation of plant populations and the persistence of selection through succession, we manipulated herbivory in a long‐term field experiment. We suppressed insects in half of 16 plots over nine years and examined the genotypic structure and chemical defense of common dandelion (Taraxacum officinale), a naturally colonizing perennial apomictic plant. Insect suppression doubled dandelion abundance in the first few years, but had negligible effects thereafter. Using microsatellite DNA markers, we genotyped >2500 plants and demonstrate that insect suppression altered the genotypic composition of plots in both sampling years. Phenotypic and genotypic estimates of defensive terpenes and phenolics from the field plots allowed us to infer phenotypic plasticity and the response of dandelion populations to insect‐mediated natural selection. The effects of insect suppression on plant chemistry were, indeed, driven both by plasticity and plant genotypic identity. In particular, di‐phenolic inositol esters were more abundant in plots exposed to herbivory (due to the genotypic composition of the plots) and were also induced in response to herbivory. This field experiment thus demonstrates evolutionary sorting of plant genotypes in response to insect herbivores that was in same direction as the plastic defensive response within genotypes.     

Herbivore induced plant volatiles: Their role in plant defense for pest management

Plants respond to herbivory through different defensive mechanisms. The induction of volatile emission is one of the important and immediate response of plants to herbivory. Herbivore-induced plant volatiles (HIPVs) are involved in plant communication with natural enemies of the insect herbivores, neighboring plants, and different parts of the damaged plant. Release of a wide variety of HIPVs in response to herbivore damage and their role in plant-plant, plant-carnivore and intraplant communications represents a new facet of the complex interactions among different trophic levels. HIPVs are released from leaves, flowers, and fruits into the atmosphere or into the soil from roots in response to herbivore attack. Moreover, HIPVs act as feeding and/or oviposition deterrents to insect pests. HIPVs also mediate the interactions between the plants and the microorganisms. This review presents an overview of HIPVs emitted by plants, their role in plant defense against herbivores and their implications for pest management.     

昆虫植食对植物有性生殖的影响

植物在个体发育的各个阶段都与不同的群落成员相互作用,如竞争的植物、有益的传粉者和敌对的植食动物。昆虫植食在各类生态系统中普遍存在,并可能对植物有性生殖产生各种影响。植食昆虫可通过对植物有性生殖结构的消耗直接对植物生殖产生影响,也可通过影响植物资源分配和花性状等改变传粉者服务,从而间接对植物有性生殖带来正面、负面或中性的影响。同一植物的植食昆虫和传粉者往往对植物的吸引性状 （如花大小、气味、颜色等）有相同的偏好,因此植食者与传粉者均能对植物有性生殖性状施加选择压力。本文从昆虫植食对植物有性生殖的直接影响、间接影响以及植食昆虫对植物有性生殖性状选择的影响三个方面进行综述,以期为昆虫植食和生物资源多样性保护相关研究提供参考。     

Leaf trichome responses to herbivory in willows: induction, relaxation and costs

Direct measurement of the carbon (C) 'cost' of mycorrhizas is problematic. Although estimates have been made for arbuscular and ectomycorrhizal symbioses, these are based on incomplete budgets or indirect measurements. Furthermore, the conventional model of unidirectional plant-to-fungus C flux is too simplistic. Net fungus-to-plant C transfer supports seedling establishment in c. 10% of plant species, including most orchids, and bidirectional C flows occur in ectomycorrhiza utilizing soil amino acids. Here, the C cost of mycorrhizas to the green orchid Goodyera repens was determined by measurement of simultaneous bidirectional fluxes of 14C labelled sources using a monoxenic system with the fungus Ceratobasidium cornigerum. Transfer of C from fungus to plant ('up-flow') occurs in the photosynthesizing orchid G. repens (max. 0.06 microg) whereas over five times more current assimilate (min. 0.355 microg) is simultaneously allocated in the reverse direction to the mycorrhizal fungus ('down-flow') after 8 d. Carbon is transferred rapidly, being detected in plant-fungal respiration within 31 h of labelling. This study provides the most complete C budget for an orchid-mycorrhizal symbiosis, and clearly shows net plant-to-fungus C flux. The rapidity of bidirectional C flux is indicative of dynamic transfer at an interfacial apoplast as opposed to reliance on digestion of fungal pelotons.     

虫害诱导的植物挥发物:植物与植食性昆虫及其天敌相互作用的进化产物

综述了植物、植食性昆虫及其天敌相互作用的进化过程。虫害诱导的植物挥发物的特征和功能是植物-植食性昆虫-天敌之间长期进化的结果。在植物、植食性昆虫与天敌相互作用的进化过程中,3个不同营养级,包括植物、植食性昆虫和天敌有着各自的调节和利用虫害诱导的植物挥发物的策略。但有一些问题,如通过实验研究得出的诱导防御在田间是否真正能起到保护作用等需进一步研究、阐明。     

Plant genotype and induced responses affect resistance to herbivores on evening primrose (Oenothera biennis)

Abstract. 1. Although both genotype and induced responses affect a plant's resistance to herbivores, little is known about their relative and interactive effects. This study examined how plant genotype of a native plant (Oenothera biennis) and induced plant responses to herbivory affect resistance to, and interactions among, several herbivores. 2. In a field experiment, genetic and environmental variation among habitats led to variation in the amount of early season damage and plant quality. The pattern of variation in early season infestation by spittlebugs (Philaenus spumarius, a piercing–sucking herbivore) negatively correlated with oviposition preference by a later feeding specialist weevil (Tyloderma foveolatum, a leaf‐chewer). 3. To determine if plant genotype and induced responses to herbivory might be responsible for these field patterns, we performed no‐choice and choice bioassays using four genotypes of O. biennis that varied in resistance. Plants were induced by either spittlebugs or weevils and assays measured the responses of the same specialist weevil as well as a generalist caterpillar (Spodoptera exigua). 4. Resistance to adult weevils was largely unaffected by plant genotype, while they experienced induced resistance following damage by conspecific weevils in no‐choice assays. Caterpillars were more strongly affected by plant genotype than induced responses in both no‐choice and choice assays, but they also fed less and experienced higher mortality on plants previously damaged by weevils. In contrast to the pattern suggested by the field experiment, spittlebugs did consistently induce resistance against either weevils or caterpillars in the bioassay experiment. 5. These results support recent findings that show herbivore species can compete via induced plant responses. Additionally, a quantitative review of the literature demonstrates that plant genotype tends to be more important than interspecific competition among herbivores (plant‐mediated or otherwise) in affecting herbivore preference and performance.     

Testing the potential for conflicting selection on floral chemical traits by pollinators and herbivores: predictions and case study

1.  There are myriad ways in which pollinators and herbivores can interact via the evolutionary and behavioural responses of their host plants.
2.  Given that both herbivores and pollinators consume and are dependent upon plant-derived nutrients and secondary metabolites, and utilize plant signals, plant chemistry should be one of the major factors mediating these interactions.
3.  Here we build upon a conceptual framework for understanding plant-mediated interactions of pollinators and herbivores. We focus on plant chemistry, in particular plant volatiles and aim to unify hypotheses for plant defence and pollination. We make predictions for the evolutionary outcomes of these interactions by hypothesizing that conflicting selection pressures from herbivores and pollinators arise from the constraints imposed by plant chemistry.
4.  We further hypothesize that plants could avoid conflicts between pollinator attraction and herbivore defence through tissue-specific regulation of pollinator reward chemistry, as well as herbivore-induced changes in flower chemistry and morphology.
5.  Finally, we test aspects of our predictions in a case study using a wild tomato species, Solanum peruvianum , to illustrate the diversity of tissue-specific and herbivore-induced differences in plant chemistry that could influence herbivore and pollinator behaviour, and plant fitness.     

Adaptations to water gradient in three Rorippa plant species correspond with plant resistance against insect herbivory under drought and waterlogged conditions

1. Plants live in environments where they are constantly, and often simultaneously, exposed to different types of biotic and abiotic stress, such as insect herbivory and water availability. How plants are adapted to abiotic conditions may determine how a surplus or shortage of water affects plant resistance to insect herbivory. Moreover, this effect may vary depending on the feeding mode of the herbivore.
2. We explored how three closely related Rorippa plant species that vary in adaptations to different water levels, resist herbivory by four different insects (aphids: Myzus persicae, Lipaphis erysimi, and caterpillars: Pieris brassicae, Plutella xylostella) under waterlogging or drought conditions. We hypothesized that plants that are differently adapted to water availability will be disparately affected by water availability in their resistance to insect herbivory.
3. On the semi-aquatic plant species Rorippa amphibia, both aphid species reached a larger colony size under drought conditions. This indicates that R. amphibia was compromised in resistance to aphid feeding when under drought conditions, to which it is less well adapted. Water conditions did not affect aphid performance on the flood-plain species Rorippa palustris. On the terrestrial plant species Rorippa sylvestris, aphids performed worse on waterlogged than drought-treated plants. Neither caterpillar species was significantly affected by the water availability of their food plant.
4. Our findings suggest that water availability can have distinct effects on plant–insect interactions. We propose that plant adaptations to water conditions can be a major predictor towards explaining the variation of effects that water availability can have on plant–insect interactions.

     

Tree seedling responses to leaf-cutting ants herbivory in Atlantic Forest restoration sites

Leaf-cutting ants (LCA) are generalist herbivores capable of causing severe plant damage. Negative impacts of ant herbivory vary according to the density of nests and availability of palatable plants; however, it is not yet clear how these herbivores affect tropical forest restoration sites. To investigate how LCA preference affects plant species performance, we evaluated the herbivory of Atta sexdens rubropilosa on native tree species seedlings in Atlantic Forest restoration sites. We expected pioneer species to suffer higher herbivory by LCA when compared with non-pioneer species and that species with higher damage will have poorer growth and higher mortality. The experiment was conducted in three restoration sites in northern Paraná state, southern Brazil, with 1,500 seedlings of 5 pioneer and 5 non-pioneer species. Sites share similar age, stand size, tree species composition, and LCA nest density. The number of attacks, degree of leaf damage, number of leaves, plant height, and survival were recorded. Specific leaf area, leaf polyphenols, flavonoids, tannins, and nitrogen content were analyzed for each species. Plant damage was similar between pioneer and non-pioneer plant species. This could be explained by trait variability among species in each group and by LCA generalist foraging. Preferred species suffered decreases in growth and survival. Less preferred species suffered fewer ant attacks and no change in performance. Results suggest that ant herbivory can influence plant species establishment and thus species composition in restoration sites by reducing performance and increasing mortality of some, but not all species, making LCA an important ecological filter.     

On the study of plant defence and herbivory using comparative approaches: how important are secondary plant compounds

Species comparisons are a cornerstone of biology and there is a long tradition of using the comparative framework to study the ecology and evolution of plant defensive traits. Early comparative studies led to the hypothesis that plant chemistry plays a central role in plant defence, and the evolution of plant secondary chemistry in response to insect herbivory remains a classic example of coevolution. However, recent comparative work has disagreed with this paradigm, reporting little connection between plant secondary chemicals and herbivory across distantly related plant taxa. One conclusion of this new work is that the importance of secondary chemistry in plant defence may have been generally overstated in earlier research. Here, we attempt to reconcile these contradicting viewpoints on the role of plant chemistry in defence by critically evaluating the use and interpretation of species correlations as a means to study defence–herbivory relationships. We conclude that the notion that plant primary metabolites (e.g. leaf nitrogen content) are the principal determinants of herbivory (or the target of natural selection by herbivores) is not likely to be correct. Despite the inference of recent community‐wide studies of herbivory, strong evidence remains for a prime role of secondary compounds in plant defence against herbivores.     

硅对植物抗虫性的影响及其机制

硅不是植物必需营养元素,但硅在提高植物对一系列非生物和生物胁迫的抗性方面都具有重要作用。综述了硅对植物抗虫性的影响及其机制。在多数植物中,增施硅肥可增强其抗虫性;所增强的抗性与硅肥种类和施用方式之间存在关系。植物组织中沉积的硅可增加其硬度和耐磨度,降低植物可消化性,从而增强植物组成性防御,包括延缓昆虫生长发育、降低繁殖力、减轻植物受害程度;植物体内的硅含量以及硅沉积的位点和排列方式影响组成性防御作用的强度。此外,硅可以调节植物诱导性防御,包括直接防御和间接防御,直接防御涉及增加有毒物质含量、产生局部过敏反应或系统获得抗性、产生有毒化合物和防御蛋白,从而延缓昆虫发育;间接防御主要通过释放挥发性化合物吸引植食性昆虫的捕食性和寄生性天敌而导致植食性昆虫种群下降。     

Induced response to herbivory in stinging hair traits of Japanese nettle (Urtica thunbergiana) seedlings in two subpopulations with different browsing pressures by sika deer

Evolutionary theories predict that natural selection favors inducible defense when the risk of predation is unpredictable. In this context, the magnitude of the induced defense in populations experiencing intermittent herbivory is predicted to be larger than that in populations experiencing constant herbivory when there is genetic differentiation between populations. To test this prediction, we conducted a clipping experiment to investigate induced response to shoot damage by the stinging hair traits of Japanese nettle (Urtica thunbergiana) seedlings. For this purpose, we studied two nettle subpopulations, one under constant browsing and another under intermittent browsing by sika deer in Nara Park, central Japan. The clipping experiment demonstrates that both subpopulations exhibited induced defenses in response to the clipping of the shoot apex as the number and length of stinging hairs increased after clipping. The subpopulation experiencing intermittent browsing exhibited smaller trait values and larger induced defenses, indicated by the number of stinging hairs on the upper leaf surface and the length of stinging hairs on both leaf surfaces compared with the subpopulation experiencing constant browsing. These results are consistent with the prediction and suggest that genetic differentiation of the induced defense between subpopulations is caused by adaptation to the herbivory regime. We discuss other plausible factors affecting the magnitude of the induced defense of the nettle subpopulations.     

Elevational gradients in plant defences and insect herbivory: recent advances in the field and prospects for future research

Classic research on elevational gradients in plant–herbivore interactions holds that insect herbivore pressure is stronger under warmer, less seasonal climates characteristic of low elevations, and that this in turn selects for increased defence in low‐ (relative to high‐) elevation plants. However, recent work has questioned this paradigm, arguing that it overly simplifies the ecological complexity in which plant–insect herbivore interactions are embedded along elevational gradients. Numerous biotic and abiotic factors vary with elevation, and their simultaneous influences are the focus of current work on elevational gradients in insect herbivory and plant defences. The present review 1) synthesizes current knowledge on elevational gradients in plant–insect herbivore interactions; 2) critically analyses research gaps and highlights recent advances that contribute to filling these gaps; and 3) outlines new research opportunities to uncover underlying mechanisms and build towards a unified theory on elevational gradients. We conclude that the next generation of studies should embrace community complexity – including multi‐trophic dynamics and the multivariate nature of plant defence – and to do so by combining observational data, manipulative experiments and emerging analytical tools.     

Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.     

Interacting effects of herbivory and fertility on a synthesized plant community

1 Outdoor microcosms were used to investigate the effects of invertebrate herbivory on plant community composition, and thereby infer possible effects on the rate of secondary succession, at differing levels of soil fertility.
2 A mixture containing 24 grassland plant species of widely different functional types was established, with 12 microcosms at each of three fertility levels. Four generalist herbivores ( Helix aspersa , Cepaea hortensis , Arianta arbustorum and Sitobion avenae ) were added to half of the microcosms. Above-ground biomass of each species was harvested after 2 years. Reproductive variables were also measured for one species, Poa annua .
3 At both moderate and high soil fertility generalist invertebrate herbivores fed selectively on early successional, fast-growing species, thus increasing the relative abundance of later successional, slow-growing species. This supports the hypothesis that herbivory increases the rate of secondary succession. Flowering and viable seed production of early successional ephemerals was also reduced by the invertebrate herbivores across a wide range of soil fertility. This would seriously reduce the ability of a species to persist in the community, thereby further hastening the rate of succession.