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1.
1. Herbivores sometimes suppress plant defences. This study tested whether the presence of pea aphids (Acyrthosiphon pisum Harris) on broad bean (Vicia faba) led to decreased secretion of extrafloral nectar (EFN) which functions as an indirect plant defence against herbivores. 2. To determine effects of aphid infestation on EFN secretion, a comparison was done between EFN secretion in uninfested plants and that in plants infested by A. pisum and another aphid species (Aphis craccivora Koch). 3. When broad bean plants were infested by A. pisum, they secreted significantly smaller amounts of EFN than did uninfested plants and A. craccivora‐infested plants. There was no significant difference in EFN secretion between uninfested plants and A. craccivora‐infested plants. The number of extrafloral nectaries did not differ among the three treatments. 4. These results suggest that A. pisum reduced EFN production in broad bean plants.  相似文献   

2.
Radhika V  Kost C  Bartram S  Heil M  Boland W 《Planta》2008,228(3):449-457
Many plants respond to herbivory with an increased production of extrafloral nectar (EFN) and/or volatile organic compounds (VOCs) to attract predatory arthropods as an indirect defensive strategy. In this study, we tested whether these two indirect defences fit the optimal defence hypothesis (ODH), which predicts the within-plant allocation of anti-herbivore defences according to trade-offs between growth and defence. Using jasmonic acid-induced plants of Phaseolus lunatus and Ricinus communis, we tested whether the within-plant distribution pattern of these two indirect defences reflects the fitness value of the respective plant parts. Furthermore, we quantified photosynthetic rates and followed the within-plant transport of assimilates with (13)C labelling experiments. EFN secretion and VOC emission were highest in younger leaves. Moreover, the photosynthetic rate increased with leaf age, and pulse-labelling experiments suggested transport of carbon to younger leaves. Our results demonstrate that the ODH can explain the within-plant allocation pattern of both indirect defences studied.  相似文献   

3.
Indirect defence via tritrophic interactions   总被引:12,自引:0,他引:12  
Many plants interact with carnivores as an indirect defence against herbivores. The release of volatile organic compounds (VOCs) and the secretion of extrafloral nectar (EFN) are induced by insect feeding, a response that is mediated by the plant hormone, jasmonic acid. Although VOCs mainly attract predatory mites and parasitic wasps, while EFN mainly attracts ants, many more animal-plant interactions are influenced by these two traits. Other traits involved in defensive tritrophic interactions are cellular food bodies and domatia, which serve the nutrition and housing of predators. They are not known to respond to herbivory, while food body production can be induced by the presence of the mutualists. Interactions among the different defensive traits, and between them and other biotic and abiotic factors exist on the genetic, physiological, and ecological levels, but so far remain understudied. Indirect defences are increasingly being discussed as an environmentally-friendly crop protection strategy, but much more knowledge on their fitness effects under certain environmental conditions is required before we can understand their ecological and evolutionary relevance, and before tritrophic interactions can serve as a reliable tool in agronomy.  相似文献   

4.
Choh Y  Kugimiya S  Takabayashi J 《Oecologia》2006,147(3):455-460
We found that intact lima bean plants increased the secretion of extrafloral nectar (EFN) after exposure to Tetranychus urticae-induced plant volatiles. Predatory mites, Phytoseiulus persimilis, dispersed more slowly from an exposed intact plant than from a control plant (plant exposed to volatiles from intact conspecific). The predators also dispersed more slowly from those plants that were provided with extra EFN than from untreated plants. We further show that EFN was a potential alternative food source for P. persimilis. From these results, we concluded that increased EFN was involved in the slow dispersal of P. persimilis from the plants exposed to herbivore-induced plant volatiles. Our data suggest that the increase of EFN in an HIPV-exposed intact plant could be an induced indirect defense against spider mites.  相似文献   

5.
Conspicuous extra-floral nectaries are inducible in Vicia faba   总被引:2,自引:0,他引:2  
Mutualistic interactions are dynamic associations that vary depending on the costs and benefits to each of the interacting parties. Phenotypic plasticity in mutualistic interactions allows organisms to produce rewards to attract mutualists when the benefits of their presence outweigh the costs of producing the rewards. In ant–plant defensive mutualisms, defences are indirect as plants produce extra‐floral nectaries (EFN) to attract predatory ants to deter herbivores. Here we demonstrate that in broad bean, Vicia faba, the overall number of EFNs on a plant increases dramatically following leaf damage. In two damage treatments, removal of: (1) one‐third of one leaf in a single leaf pair or (2) one‐third of both halves of a single leaf pair, resulted in a 59 and 106% increase in the number of EFNs on the plants, respectively, over 1 week. We suggest that the increased production of visually conspicuous EFNs is an adaptive inducible response, to attract predatory arthropods when risk of herbivory increases.  相似文献   

6.
Plants in more than 300 genera produce extrafloral nectar (EFN) to attract carnivores as a means of indirect defence against herbivores. As EFN is secreted at nectaries that are not physically protected from the environment, and contains carbohydrates and amino acids, EFN must be protected from infestation by micro-organisms. We investigated the proteins and anti-microbial activity in the EFN of two Central American Acacia myrmecophytes ( A. cornigera and A. hindsii ) and two related non-myrmecophytes ( A. farnesiana and Prosopis juliflora ). Acacia myrmecophytes secrete EFN constitutively at high rates to nourish the ants inhabiting these plants as symbiotic mutualists, while non-myrmecophytes secrete EFN only in response to herbivore damage to attract non-symbiotic ants. Thus, the quality and anti-microbial protection of the EFN secreted by these two types of plants were likely to differ. Indeed, myrmecophyte EFN contained significantly more proteins than the EFN of non-myrmecophytes, and was protected effectively from microbial infestation. We found activity for three classes of pathogenesis-related (PR) enzymes: chitinase, β-1,3-glucanase and peroxidase. Chitinases and β-1,3-glucanases were significantly more active in myrmecophyte EFN, and chitinase at the concentrations found in myrmecophyte EFN significantly inhibited yeast growth. Of the 52 proteins found in A. cornigera EFN, 28 were annotated using nanoLC-MS/MS data, indicating that chitinases and glucanases contribute more than 50% of the total protein content in the EFN of this myrmecophyte. Our study demonstrates that PR enzymes play an important role in protecting EFN from microbial infestation.  相似文献   

7.
Background and Aims Many plants produce extrafloral nectar (EFN), and increase production following above-ground herbivory, presumably to attract natural enemies of the herbivores. Below-ground herbivores, alone or in combination with those above ground, may also alter EFN production depending on the specificity of this defence response and the interactions among herbivores mediated through plant defences. To date, however, a lack of manipulative experiments investigating EFN production induced by above- and below-ground herbivory has limited our understanding of how below-ground herbivory mediates indirect plant defences to affect above-ground herbivores and their natural enemies.Methods In a greenhouse experiment, seedlings of tallow tree (Triadica sebifera) were subjected to herbivory by a specialist flea beetle (Bikasha collaris) that naturally co-occurs as foliage-feeding adults and root-feeding larvae. Seedlings were subjected to above-ground adults and/or below-ground larvae herbivory, and EFN production was monitored.Key Results Above- and/or below-ground herbivory significantly increased the percentage of leaves with active nectaries, the volume of EFN and the mass of soluble solids within the nectar. Simultaneous above- and below-ground herbivory induced a higher volume of EFN and mass of soluble solids than below-ground herbivory alone, but highest EFN production was induced by above-ground herbivory when below-ground herbivores were absent.Conclusions The induction of EFN production by below-ground damage suggests that systemic induction underlies some of the EFN response. The strong induction by above-ground herbivory in the absence of below-ground herbivory points to specific induction based on above- and below-ground signals that may be adaptive for this above-ground indirect defence.  相似文献   

8.
Plants show defensive responses after exposure to volatiles from neighbouring plants infested by herbivores. When a plant’s neighbours host only species of herbivores that do not feed on the plant itself, the plant can conserve energy by maintaining a low defence level. An intriguing question is whether plants respond differently to volatiles from plants infested by herbivores that pose greater or lesser degrees of danger. We examined the secretion of extrafloral nectar (EFN) in lima bean plants exposed to volatiles from cabbage plants infested by common cutworm, two-spotted spider mites, or diamondback moth larvae. Although the first two herbivore species feed on lima bean plants, diamondback moth larvae do not. As a control, lima bean plants were exposed to volatiles from uninfested cabbage plants. Only when exposed to volatiles from cabbage plants infested by spider mites did lima bean plants significantly increase their EFN secretion compared with the control. Increased EFN secretion can function as an indirect defence by supplying the natural enemies of herbivores with an alternative food source. Of the three herbivore species, spider mites were the most likely to move from cabbage plants to lima bean plants and presumably posed the greatest threat. Although chemical analyses showed differences among treatments in volatiles produced by herbivore-infested cabbage plants, which compounds or blends triggered the increased secretion of EFN by lima bean plants remains unclear. Thus, our results show that plants may tune their defence levels according to herbivore risk level.  相似文献   

9.
Extrafloral nectar (EFN) plays an important role as plant indirect defence through the attraction of defending ants. Like all rewards produced in the context of a mutualism, however, EFN is in danger of being exploited by non-ant consumers that do not defend the plant against herbivores. Here we asked whether plants, by investing more in EFN, can improve their indirect defence, or rather increase the risk of losing this investment to EFN thieves. We used the obligate plant-ant Acacia-Pseudomyrmex system and examined experimentally in the field during the dry and the rainy seasons how variations in EFN secretion are related to (i) ant activity, to (ii) the ant-mediated defence against herbivores and (iii) the exploitation of EFN by non-ant consumers. Extrafloral investment enhanced ant recruitment and was positively related to the ant mediated defence against herbivores. The ant-mediated protection from exploiters also increased in proportion to the nectar sugar concentration. Although the daily peak of EFN production coincided with the highest activity of EFN thieves, Pseudomyrmex ferrugineus ants protected this resource effectively from exploiters. Nevertheless, the defensive effects by ants differed among seasons. During the dry season, plants grew slower and secreted more EFN than in the rainy season, and thus, experienced a higher level of ant-mediated indirect defence. Our results show that an increased plant investment in an indirect defence trait can improve the resulting defensive service against both herbivores and exploiters. EFN secretion by obligate ant-plants represents a defensive trait for which the level of investment correlates positively with the beneficial effects obtained.  相似文献   

10.
1. Predatory ants may reduce infestation by herbivorous insects, and slow‐moving Lepidopteran larvae are often vulnerable on foliage. We investigate whether caterpillars with morphological or behavioural defences have decreased risk of falling prey to ants, and if defence traits mediate host plant use in ant‐rich cerrado savanna. 2. Caterpillars were surveyed in four cerrado localities in southeast Brazil (70–460 km apart). The efficacy of caterpillar defensive traits against predation by two common ant species (Camponotus crassus, C. renggeri) was assessed through experimental trials using caterpillars of different species and captive ant colonies. 3. Although ant presence can reduce caterpillar infestation, the ants' predatory effects depend on caterpillar defence traits. Shelter construction and morphological defences can prevent ant attacks (primary defence), but once exposed or discovered by ants, caterpillars rely on their size and/or behaviour to survive (secondary defence). 4. Defence efficiency depends on ant identity: C. renggeri was more aggressive and lethal to caterpillars than C. crassus. Caterpillars without morphological defences or inside open shelters were found on plants with decreased ant numbers. No unsheltered caterpillar was found on plants with extrafloral nectaries (EFNs). Caterpillars using EFN‐bearing plants lived in closed shelters or presented morphological defences (hairs, spines), and were less frequently attacked by ants during trials. 5. The efficiency of defences against ants is thus crucial for caterpillar survival and determines host plant use by lepidopterans in cerrado. Our study highlights the effect of EFN‐mediated ant‐plant interactions on host plant use by insect herbivores, emphasizing the importance of a tritrophic viewpoint in risky environments.  相似文献   

11.
Tomato plants release volatile organic compounds (VOCs) following insect or mechanical damage. In this study, the constitutive and wound-induced emission levels of VOCs in suppressor of prosystemin-mediated responses2 (spr2) mutant plants, compromised in linolenic acid (LA) and jasmonic acid (JA) synthesis, and in 35S::prosystemin (35S::prosys) plants, having upregulated direct defence responses, were compared. The spr2 mutants produced constitutively lower levels of VOCs, which were nonetheless increased in response to (a)biotic damage, although at lower levels than wild-type (Wt) and 35S::prosys plants. No significant differences in VOC emissions were detected between the latter two genotypes, thereby suggesting that systemin does not regulate indirect defence responses, whereas differences in fatty acid composition in spr2 plants led to the predominant emission of saturated C6 volatiles in response to wounding. The expression of 1-deoxy-D-xylulose 5-phosphate synthase (DXS2), a key gene involved in VOC synthesis in the chloroplast, was only upregulated in Manduca sexta L.-damaged Wt and 35S::prosys plants. However, its expression was restored in spr2 plants by exogenous LA or JA, suggesting that abated VOC emissions in spr2 plants are correlated with lowered DXS2 expression. Bioassays with two different insects showed that adult females significantly preferred spr2 plants, indicating that lowered levels of VOCs in tomato influence plant selection by insects during oviposition.  相似文献   

12.
Extrafloral nectar (EFN) mediates food‐for‐protection mutualisms between plants and insects and provides plants with a form of indirect defense against herbivory. Understanding sources of variation in EFN production is important because such variations affect the number and identity of insect visitors and the effectiveness of plant defense. Light represents a potentially crucial tool for regulating resource allocation to defense, as it not only contributes energy but may help plants to anticipate future conditions. Low red/far‐red (R/FR) light ratios can act as a signal of the proximity of competing plants. Exposure to such light ratios has been shown to promote competitive behaviors that coincide with reduced resource allocation to direct chemical defenses. Little is known, however, about how such informational light signals might affect indirect defenses such as EFN, and the interactions that they mediate. Through controlled glasshouse experiments, we investigated the effects of light intensity, and R/FR light ratios, on EFN production in Senna mexicana var. chapmanii. Plants in light‐limited conditions produced significantly less EFN, and leaf damage elicited increased EFN production regardless of light conditions. Ratios of R/FR light, however, did not appear to affect EFN production in either damaged or undamaged plants. Understanding the effects of light on indirect defenses is of particular importance for plants in the threatened pine rockland habitats of south Florida, where light conditions are changing in predictable ways following extensive fragmentation and subsequent mismanagement. Around 27% of species in these habitats produce EFN and may rely on insect communities for defense.  相似文献   

13.
Folivores are major plant antagonists in most terrestrial ecosystems. However, the quantitative effects of leaf area loss on multiple interacting plant traits are still little understood. We sought to contribute to filling this lack of understanding by applying different types of leaf area removal (complete leaflets versus leaflet parts) and degrees of leaf damage (0, 33 and 66%) to lima bean (Phaseolus lunatus) plants. We quantified various growth and fitness parameters including above‐ and belowground biomass as well as the production of reproductive structures (fruits, seeds). In addition, we measured plant cyanogenic potential (HCNp; direct chemical defence) and production of extrafloral nectar (EFN; indirect defence). Leaf damage reduced above‐ and belowground biomass production in general, but neither variation in quantity nor type of damage resulted in different biomass. Similarly, the number of fruits and seeds was significantly reduced in all damaged plants without significant differences between treatment groups. Seed mass, however, was affected by both type and quantity of leaf damage. Leaf area loss had no impact on HCNp, whereas production of EFN decreased with increasing damage. While EFN production was quantitatively affected by leaf area removal, the type of damage had no effect. Our study provides a thorough analysis of the quantitative and qualitative effects of defoliation on multiple productivity‐related and defensive plant traits and shows strong differences in plant response depending on trait. Quantifying such plant responses is vital to our understanding of the impact of herbivory on plant fitness and productivity in natural and agricultural ecosystems.  相似文献   

14.
Assessment of host plant specificity is a critical step in the evaluation of classical biological control agents of weeds which is necessary for avoiding possible damage to non-target plants. Volatile organic compounds (VOCs) emitted by plants likely play an important role in determining which plants attract and are accepted by a prospective arthropod agent. However, current methods to evaluate host plant specificity usually rely on empirical choice and no-choice behavioural experiments, with little knowledge about what chemical or physical attributes are stimulating the insect. We conducted experiments to measure the quantitative and qualitative effects on emission of VOCs caused by simple mechanical damage to leaves of plants known to differ in suitability and attractiveness to a prospective agent. More VOCs were detected from damaged than from undamaged leaves for all three species tested. Discriminant analysis was able to correctly distinguish the taxonomic identity of all plants based on their VOC profiles; however, the VOCs that discriminated species among undamaged leaves were completely different from those that discriminated among damaged leaves. Thus, damaged and undamaged plants present different VOC profiles to insects, which should be considered when conducting host plant specificity experiments. An unacceptable non-target plant, Centaurea cineraria, emitted all except one of the VOCs that were emitted by its preferred host plant, Centaurea solstitialis, indicating the importance of compounds that are repellant in host plant specificity. Centaurea cyanus emitted fewer VOCs than C. solstitialis, which suggests that it lacked some VOCs important for host plant recognition.  相似文献   

15.
Much effort has been devoted to understanding the function of extrafloral nectaries (EFNs) for ant–plant–herbivore interactions. However, the pattern of evolution of such structures throughout the history of plant lineages remains unexplored. In this study, we used empirical knowledge on plant defences mediated by ants as a theoretical framework to test specific hypotheses about the adaptive role of EFNs during plant evolution. Emphasis was given to different processes (neutral or adaptive) and factors (habitat change and trade‐offs with new trichomes) that may have affected the evolution of ant–plant associations. We measured seven EFN quantitative traits in all 105 species included in a well‐supported phylogeny of the tribe Bignonieae (Bignoniaceae) and collected field data on ant–EFN interactions in 32 species. We identified a positive association between ant visitation (a surrogate of ant guarding) and the abundance of EFNs in vegetative plant parts and rejected the hypothesis of phylogenetic conservatism of EFNs, with most traits presenting K‐values < 1. Modelling the evolution of EFN traits using maximum likelihood approaches further suggested adaptive evolution, with static‐optimum models showing a better fit than purely drift models. In addition, the abundance of EFNs was associated with habitat shifts (with a decrease in the abundance of EFNs from forest to savannas), and a potential trade‐off was detected between the abundance of EFNs and estipitate glandular trichomes (i.e. trichomes with sticky secretion). These evolutionary associations suggest divergent selection between species as well as explains K‐values < 1. Experimental studies with multiple lineages of forest and savanna taxa may improve our understanding of the role of nectaries in plants. Overall, our results suggest that the evolution of EFNs was likely associated with the adaptive process which probably played an important role in the diversification of this plant group.  相似文献   

16.
Galen C 《Oecologia》2005,144(1):80-87
According to the distraction hypothesis, extrafloral nectaries (EFN) evolved under selection to entice ants away from floral nectaries, reducing ant-mediated damage to flowers and/or interference with pollinators. Predator-satiation, through production of nectar in either surplus flowers or EFN, provides an alternative mechanism for reducing the impact of ants as flower visitors. I tested these two hypotheses by experimentally adding EFN to flowering plants of the alpine wildflower, Polemonium viscosum, and by surveying the relationship between ant visitation and nectary number in nature. Plants of P. viscosum lack EFN and experience flower damage by ants of Formica neorufibarbus gelida. Ant behavior was compared on plants with five flowers and three experimental EFN and on controls with equal floral display, but no EFN. Addition of EFN increased flower visitation by ants. The effect of EFN on flower visitation did not depend on proximity of EFN to flowers or attractiveness of EFN to ants. Findings suggest that ants perceived patch quality on a whole plant basis, rather than responding to EFN and flowers as distinct nectar patches. Ant visitation did not keep pace with nectary number in nature. The relationship between ant visitation and nectary number per plant was weak and shallow as predicted under satiation. Ant foraging choices on experimental inflorescences showed that ants bypass flowers avoided by earlier ants, enhancing probability of escape via satiation. Results do not support the idea that EFN evolve to reduce flower visitation by ants, but show instead that nectar in surplus flowers can satiate ants and reduce their negative impacts on flower function and integrity.  相似文献   

17.
Some volatile organic chemicals (VOCs), such as terpenes, are responsible for communication between plants. We assessed the priming of defense responses in lima bean by exposing the plants to transgenic-plant-volatiles [(E)-β-ocimene] emitted from transgenic tobacco plants (NtOS2). As it was previously shown that the first receiver lima bean plants, which were infested with spider mites after having been exposed to (E)-β-ocimene from NtOS2, were highly induced to emit VOCs, we analyzed the VOCs emitted from a second set of receiver plants (second receiver plants) exposed to the infested, first receiver plants. In response to feeding by spider mites, two homoterpenes [(E)-4,8-dimethyl-1,3,7-nonatriene and (E,E)-4,8,12-trimethyltrideca-1,3,7,11-tetraene] were more highly emitted from the second receiver plants in response to spider mite attack, in comparison to the levels emitted from plants that had been placed near infested, wild-type (WT)-volatile-exposed plants. These data suggest that transgenic-plant-volatile-mediated, multiple-plant communication can function in plant defenses.  相似文献   

18.
While nectaries are commonly found in flowers, some plants also form extrafloral nectaries on stems or leaves. For the first time in the family Brassicaceae, here we report extrafloral nectaries in Brassica juncea. The extrafloral nectar (EFN) was secreted from previously amorphic sites on stems, flowering stalks and leaf axils from the onset of flowering until silique formation. Transverse sections at the point of nectar secretion revealed a pocket‐like structure whose opening was surrounded by modified stomatal guard cells. The EFN droplets were viscous and up to 50% of the total weight was sugars, 97% of which was sucrose in the five varieties of B. juncea examined. Threonine, glutamine, arginine and glutamate were the most abundant amino acids. EFN droplets also contained glucosinolates, mainly gluconapin and sinigrin. Nectar secretion was increased when the plants were damaged by chewing above‐ and belowground herbivores and sap‐sucking aphids. Parasitoids of each herbivore species were tested for their preference, of which three parasitoids preferred EFN and sucrose solutions over water. Moreover, the survival and fecundity of parasitoids were positively affected by feeding on EFN. We conclude that EFN production in B. juncea may contribute to the indirect defence of this plant species.  相似文献   

19.
Obligate ant plants (myrmecophytes) in the genus Macaranga produce energy- and nutrient-rich food bodies (FBs) to nourish mutualistic ants which live inside the plants. These defend their host against biotic stress caused by herbivores and pathogens. Facultative, 'myrmecophilic' interactions are based on the provision of FBs and/or extrafloral nectar (EFN) to defending insects that are attracted from the vicinity. FB production by the myrmecophyte, M. triloba, was limited by soil nutrient content under field conditions and was regulated according to the presence or absence of an ant colony. However, increased FB production promoted growth of the ant colonies living in the plants. Ant colony size is an important defensive trait and is negatively correlated to a plant's leaf damage. Similar regulatory patterns occurred in the EFN production of the myrmecophilic M. tanarius. Nectar accumulation resulting from the absence of consumers strongly decreased nectar flow, which increased again when consumers had access to the plant. EFN flow could be induced via the octadecanoid pathway. Leaf damage increased levels of endogenous jasmonic acid (JA), and both leaf damage and exogenous JA application increased EFN flow. Higher numbers of nectary visiting insects and lower numbers of herbivores were present on JA-treated plants. In the long run, this decreased leaf damage significantly. Ant food production is controlled by different regulatory mechanisms which ensure that costs are only incurred when counterbalanced by defensive effects of mutualistic insects.  相似文献   

20.
Abiotic conditions can increase the costs of services and/or the benefits of rewards provided by mutualistic partners. Consequently, in some situations, the outcome of mutualisms can move from beneficial to detrimental for at least one partner. In the case of protective mutualisms between ant bodyguards and plants bearing extrafloral nectaries (EFNs), plants from arid environments face a trade‐off between EFN production and maintenance and water and carbon economy. This trade‐off may increase EFN costs and decrease their value as a defensive strategy to plants in such environments. Despite this, the presence of EFNs is an ubiquitous trait in plants from arid environments, suggesting that they provide greater benefits to plants in these environments to compensate for their higher costs. We used a meta‐analysis to investigate if such benefits do increase with decreasing water availability and the possible underlying causes (such as ant behaviour or ant diversity). As predicted, ant effect on EFN plants performance increased as mean annual precipitation decreased. We also found that the frequency of dominant ants on EFN plants increased in drier areas. Due to the more aggressive behaviour of dominant ants, we suggest that they represent an important factor shaping the adaptive value of EFNs to plants in arid environments.  相似文献   

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