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1.
Lima bean (Phaseolus lunatus) features two inducible indirect defences to protect itself against herbivores. Besides the emission of plant volatiles, extrafloral nectar is secreted to attract carnivorous arthropods to herbivore-damaged plants. The activation of both putative defences efficiently protects Lima beans from leaf damage. In a field experiment in Mexico, we studied whether extrafloral nectar alone can benefit the Lima bean under natural conditions. An artificial blend mimicking natural nectar both qualitatively and quantitatively was repeatedly applied to Lima bean tendrils. Ants, wasps and flies were significantly more abundant on treated tendrils than on untreated controls already after 1 week (i.e. after two treatment applications). Sticky traps were used to assess the functional groups of flying insects attracted to the Lima beans. After 24 h, 71% of all trapped flies and 98% of all wasps belonged to families comprising either parasitoid or predatory species. This observation suggests that also some of the flying visitors have played a role as putative defenders of Lima beans. Most of the trapped flies belonged to the families Dolichopodidae and Phoridae (each ca. one third of all individuals). Two thirds of the wasps belonged to Chalcidoidea (68%). All ant species that had been collected manually belonged to generalist genera with Camponotus novogranadensis and Cephalotes minutus being most regularly encountered on study tendrils. An additional experiment, where both ‘nectar’ and ‘control’ tendrils were treated with artificial nectar, revealed that ants responded with an increased abundance on tendrils that had experienced the ‘nectar’ treatment before.After 25 days, the treated tendrils showed a significantly reduced herbivory as compared to controls. The mere presence of increased amounts of extrafloral nectar thus can benefit the Lima bean under natural conditions.

Zusammenfassung

Die Limabohne (Phaseolus lunatus) verfügt über zwei induzierbare, indirekte Verteidigungsformen zur Abwehr von Herbivoren. Neben der Emission volatiler Verbindungen ist die Limabohne zusätzlich dazu in der Lage, extrafloralen Nektar zu sezernieren. Beides dient der Anlockung von Fraßfeinden zu den von Herbivoren befallenen Pflanzen. In einem Freilandexperiment in Mexiko wurde untersucht, ob die Limabohne unter natürlichen Bedingungen von der Sekretion extrafloralen Nektars profitiert. Hierzu wurde ein künstliches Nektargemisch wiederholt auf Limabohnenranken aufgetragen, welches natürlichen Nektar quantitativ und qualitativ imitierte. Bereits nach einer Woche (d.h. nach zwei Behandlungen) war die Abundanz von Ameisen, Fliegen und Wespen auf behandelten Ranken signifikant höher als auf unbehandelten Kontrollranken. Zur Erfassung der zur Limabohne angelockten fliegender Insekten sowie deren Zugehörigkeit zu funktionellen Gruppen wurden die Versuchsranken mit Klebefallen bestückt. Mehr als zwei Drittel der nach 24 h gefangenen Fliegen und 98% aller Wespen gehörten parasitisch oder räuberisch lebenden Fliegen- bzw. Wespen-Familien an. Diese Beobachtung legt nahe, dass nicht nur Ameisen, sondern auch einige der gefangenen fliegenden Besucher eine Rolle als potentielle Verteidiger der Limabohne gespielt haben könnten. Von den gefangen Fliegen gehörten die meisten den Familien Dolichopodidae und Phoridae (je ca. ein Drittel aller gefangenen Individuen) an, wogegen die Chalcidoidea zwei Drittel (68%) der gefangenen Wespen ausmachten. Unter den durch Handaufsammlung gefangenen Ameisen gehörten Camponotus novogranadensis und Cephalotes minutus zu den am häufigsten auf behandelten Ranken angetroffen Arten. Ein zusätzliches Experiment, in dem das künstliche Nektargemisch sowohl auf ‘Nektar’- als auch auf ‘Kontroll’-Ranken aufgebracht wurde, ergab, dass die Ameisen mit einer erhöhten Abundanz auf solchen Ranken reagierten, die bereits vorher die, Nektar’-Behandlung erfahren hatten.Nach 25 Tagen zeigten behandelte Ranken signifikant weniger Blattfraß im Vergleich zu unbehandelten Kontrollranken. Die bloße Erhöhung der Menge an extrafloralem Nektar reichte offensichtlich dazu aus, unter natürlichen Bedingungen wachsenden Limabohnen einen Vorteil zu verschaffen.  相似文献   

2.
Laird RA  Addicott JF 《Oecologia》2007,152(3):541-551
Arbuscular mycorrhizal fungi (AMF) can alter the physiology and morphology of their host plant, and therefore may have indirect effects on insect herbivores and pollinators. We conducted this study to test the hypothesis that AMF can also affect insects involved in protection-for-food mutualisms. We examined the constitutive and inducible production of food rewards [extrafloral (EF) nectaries] in Vicia faba plants by manipulating the presence/absence of AMF and by simulating various levels of herbivory. Plants inoculated with AMF produced significantly fewer EF nectaries than uninoculated plants, even after accounting for differences in plant growth. In contrast to earlier studies, EF nectaries were not inducible: damaged plants produced significantly fewer EF nectaries than undamaged plants. Moreover, the effects of mycorrhizal and damage status on EF nectary production were additive. The reduction in EF nectaries in mycorrhizal plants potentially represents a mechanism for indirect effects of AMF on the protective insects that exploit EF nectaries as a food source (e.g., ants). Reduced reward size should result in reduced protection by ants, and could therefore be a previously unappreciated cost of the mycorrhizal symbiosis to host plants. However, the overall effect of AMF will depend upon the extent to which the reduction of EF nectaries affects the number and activity of ants and the extent to which AMF alter other aspects of host plant physiology. Our results emphasize the complexity of multitrophic interactions, particularly those that span belowground and aboveground ecology.  相似文献   

3.
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   

4.
1. Sympatric populations of insects adapted to different host plants are good model systems not only to study how they adapt to the chemistry of their food plant, but also to investigate whether morphological modifications evolved enabling them to live successfully on a certain plant species. 2. The pea aphid, Acyrthosiphon pisum (Harris) encompasses at least 11 genetically distinct sympatric host races, each showing a preference for a certain legume species. The leaflet surfaces of these legumes differ considerably in their wax coverage. 3. It was investigated whether the attachment structures of three pea aphid genotypes from different host races are adapted to the different surface properties of their host plants and whether they show differences in their attachment ability on the respective host and non‐host plants. 4. The surface morphology of plants and aphid tarsi was examined using SEM (scanning electron microscopy). The ability of the aphids to walk on specific surfaces was tested using traction force measurements. 5. The presence of wax blooms on the leaflets lowers the aphids' attachment ability considerably and diminishes their subsequent attachment on ‘neutral’ surfaces like glass. The pea aphid host races differ in their ability to walk on certain surfaces. However, the genotype from the adapted aphid host race was not necessarily the one with the best walking performance on their host plant. All aphids, regardless of the original host plant, were most efficient on the neutral control surface glass. The general host plant Vicia faba was the plant with the most favourable surface for all aphid host races.  相似文献   

5.
6.
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   

7.
Proof of the existence of aphid-derived molecules that induce emission of volatile organic compounds as indirect defensive responses in plants (hereafter called elicitors) has not yet been obtained. We showed here the presence and some aspects of the chemical nature of these elicitors. Exogenous application of the fed diet (liquid diet previously fed on by the Pea aphid, Acyrthosiphon pisum) on the damaged part of leaves of broad bean plants, made the plants more attractive to a parasitic wasp, Aphidius ervi, than application of the unfed diet. This result suggested that elicitors existed in the fed diet. Chemical characterization using ultrafiltration and subsequent bioassay revealed that the responsible factor was larger than 3 kDa. Heating of the fed diet did not inactivate its ability to induce attractive volatiles, which suggested thermotolerance of the elicitor. These results contribute to the mechanistic understanding of plant defenses against aphids.  相似文献   

8.
This study sheds light on a poorly understood area in insect-plant-microbe interactions,focusing on aphid probing and feeding behavior on plants with varying levels of arbuscular mycorrhizal(AM)fungus root colonization.It investigates a commonly occurring interaction of three species:pea aphid Acyrthosiphon pisum,barrel medic Medicago truncatula,and the AM fungus Rhizophagus irregularis,examining whether aphid-feeding behavior changes when insects feed on plants at different levels of AM fungus colonization(42% and 84% root length colonized).Aphid probing and feeding behavior was monitored throughout 8 h of recording using the electrical penetration graph(EPG)technique,also,foliar nutrient content and plant growth were measured.Summarizing,aphids took longer to reach their 1st sustained phloem ingestion on the 84% AM plants than on the 42% AM plants or on controls.Less aphids showed phloem ingestion on the 84% AM plants relative to the 42% AM plants.Shoots of the 84% AM plants had higher percent carbon(43.7%)relative to controls(40.5%),and the 84% AM plants had reduced percent nitrogen(5.3%)relative to the 42% AM plants(6%).In conclusion,EPG and foliar nutrient data support the hypothesis that modifications in plant anatomy(e.g.,thicker leaves),and poor food quality(reduced nitrogen)in the 84% AM plants contribute to reduced aphid success in locating phloem and ultimately to differences in phloem sap ingestion.This work suggests that M.truncatula plants benefit from AM symbiosis not only because of increased nutrient uptake but also because of reduced susceptibility to aphids.  相似文献   

9.
Abstract

We studied the effects of previous infestation of broad bean plants by pea aphids Acyrthosiphon pisum on the performance of conspecific nymphs on the plants and the involvement of jasmonic acid (JA)-related defenses. The time needed for newly emerged nymphs to become reproductive adults on broad bean plants previously infested by conspecifics (pre-infested plants) was significantly shorter than on uninfested (control) broad bean plants. The total numbers of nymphs produced by aphids on preinfested and control plants were not significantly different. Preinfested plants produced significantly less endogenous JA than that control plants did. To test the effect of JA decreases, we conducted experiments on the developmental duration of nymphs on broad bean plants treated with JA (JA-treated plants) before infestation. The time needed for nymphs to become reproductive adults on JA treated preinfested broad bean plants was not significantly different from that on JA-treated control plants. The results suggested a possible parental care by pea aphids: the adult aphids manipulated JA-related defenses in broad bean plants that had positive effects for their offspring.  相似文献   

10.
Acyrthosiphon pisum is a polyphagous aphid of major importance on the pea crop to which few resistant cultivars are available. In this study, we screened a germplasm collection of Pisum spp. under field conditions over two seasons yielding the identification of a number of accessions with intermediate levels of resistance. Selected accessions were further studied under semi‐controlled and controlled conditions in no choice and choice assays to validate the responses, and to further characterise the mechanisms of resistance operative. Results elucidated the valuable resistance of accession P40 (Pisum sativum ssp. sativum) and P665 (P. sativum ssp. syriacum), with the combination of both antixenosis, by reducing aphid preference, and antibiosis, by diminishing aphid proliferation.  相似文献   

11.
The effect of temperature on the life table of Acyrthosiphon pisum reared on Pisum sativum was evaluated under laboratory conditions using temperatures of 10, 15, 20, 25, 30, and 35°C. The development time of juvenile A. pisum decreased with increasing temperature (from 21.3 days at 10°C to 4.7 days at 35°C). Adult longevity also decreased with increasing temperature (from 53.2 days at 10°C to 2.3 days at 35°C). Interestingly, 70% and 25% of A. pisum nymphs reared at 30°C and 35°C, respectively, successfully developed into adults. These temperatures have previously been considered unsuitable for A. pisum development. However, adult aphids reared at 30°C and 35°C failed to reproduce. Linear regression analysis revealed that the lower development threshold of A. pisum was 153.1 degree‐days above 1.9°C. Maximal average reproductive capability was observed at 10°C for A. pisum adults, with each adult producing more than 120 nymphs. The intrinsic rate of increase (rm) of A. pisum increased from 0.124/day at 10°C to 0.337/day at 25°C, whereas opposite trends were observed for the net reproductive rate (R0) and the mean generation time (GT). At 20°C and 25°C, the intrinsic rate of increase of A. pisum was significantly higher than at 10°C and 15°C (P < 0.0001), indicating that 20°C and 25°C are within the optimal range for the growth of A. pisum, and that 30°C is beyond the upper threshold limit for reproduction, which involves a temperature range that is narrower than that of the survival range (upper limit is unknown, but above 35°C).  相似文献   

12.
1. Aphid natural enemies include not only predators and parasitoids but also pathogens, of which fungi are the most studied for biological control. While wing formation in aphids is induced by abiotic conditions, it is also affected by biotic interactions with their arthropod natural enemies. Wing induction via interactions with arthropod natural enemies is mediated by the increase in their physical contact when alarmed (pseudo‐crowding). Pathogenic fungi do not trigger this alarm behaviour in aphids and, therefore, no pseudo‐crowding occurs. 2. We hypothesise that, while pathogenic fungi will stimulate maternally induced wing formation, the mechanism is different and is influenced by pathogen specificity. We tested this hypothesis using two entomopathogenic fungi, Pandora neoaphidis and Beauveria bassiana, an aphid specialist and a generalist respectively, on the pea aphid, Acyrthosiphon pisum Harris. 3. We first demonstrate that pea aphids infected with either pathogen and maintained in groups on broad bean plants produced a higher proportion of winged morphs than uninfected control aphids. We then show that, when maintained in isolation, aphids infected with either pathogen also produced higher proportions of winged offspring than control aphids. There was no difference between P. neoaphidis and B. bassiana in their effects on wing induction in either experiment. 4. Unlike the effect of predators and parasitoids on pea aphid wing induction, the effect of pathogens is independent of physical contact with other aphids, suggesting that physiological cues induce wing formation in infected aphids. It is possible that aphids benefit from wing induction by escaping infected patches whilst pathogens may benefit through dispersion. Possible mechanisms of wing induction are discussed.  相似文献   

13.
1. Plant defence of Viburnum shrubs against oviposition by its specialist herbivore, the viburnum leaf beetle [VLB Pyrrhalta viburni(Paykull)], involves an egg‐crushing wound response in twigs. Although the response is variable among Viburnum species, it can have a strong impact on egg survivorship. Beetles typically aggregate egg masses with conspecifics along infested twigs, forming clusters that can overwhelm the twig response. It was investigated whether twig responses and beetle oviposition behaviour vary seasonally. 2. In a field experiment, twig defences decreased towards the end of the VLB oviposition period: wound response of the North American Viburnum dentatum L. and the European V. opulus L. was reduced by 100% and 54%, respectively, in September compared with the July to August period. 3. Oviposition trials demonstrated a corresponding behavioural change: VLB females displayed aggregative oviposition in August, but not in September. 4. Further tests revealed that late‐season VLB females reverted to aggregative oviposition after being kept on uninfested twigs, whereas females kept on heavily infested twigs did not. This behavioural change suggests that relaxation of aggregative oviposition originates from cues associated with high densities of egg masses. 5. Relaxation of aggregative oviposition may be adaptive (and beneficial for invasion) on shrubs with low levels of defences by reducing intra‐specific competition.  相似文献   

14.
1. Plants can induce a response when they are attacked by herbivores. Although the induction of responses by herbivory in both flowers and leaves is relatively well studied, whether florivory (feeding of flowers) can also induce responses in flowers and leaves is less well explored and there are still unanswered questions. These include whether plants exhibit different levels of induced responses depending on the length of exposure to the florivores. 2. To address this knowledge gap, this study used a tropical floriphilic katydid, Phaneroptera brevis, and its non‐native food plant, Lantana camara. Nursery experiments were performed in which dry matter content and anthocyanin concentration of the flower (corolla and stamens), leaf dry matter content, and leaf blade punch resistance were measured at three time phases before and after exposing the plant to the katydid individuals for 0–7 days. 3. It was demonstrated that increasing the length (days) of exposure to the katydid individual leads to higher levels of induced plant response (leaf blade punch resistance), but only in the leaves. It was also shown that higher levels of induced plant response owing to the increase in the length of exposure to the katydid individual was not observed beyond the first set of leaves developed after the exposure. 4. These results address the knowledge gap and show that plants can exhibit different levels of induced responses depending on the length of exposure to florivores. This study thus highlights the far‐reaching importance of florivory on plants.  相似文献   

15.
16.
In this study, we evaluate the temporal variation in extrafloral nectaries (EFNs) secretion in different ontogenic stages of Alibertia verrucosa (Rubiaceae) fruits in a Neotropical savanna. We observe greater nectar secretion rate in fruits of intermediate size compared with young or ripe fruit, indicating that they are possibly more protected by ants. In addition, the nectar secretion was higher at night, a pattern that could be associated with an increase of herbivore pressure and water stress during the daylight hours. In fact, due to the high temperature and low humidity during the day in savannas, most herbivores display strong nocturnal activity, and plants can avoid nectar secretion in this period. Our results indicate that A. verrucosa can change the ants' attraction according to EFN secretion during the ontogenic stages of the fruits, probably secreting more nectar when the biotic defense are more necessary for the protection of the fruits and the plant as a whole.  相似文献   

17.
The symbiotic bacteria Buchnera contribute to the nutrition of pea aphids, Acyrthosiphon pisum, through the provision of essential amino acids which are lacking in the diet. However, chemically defined diets, containing nutritionally adequate amounts of essential amino acids, fail to rescue aposymbiotic aphids, in which the bacteria have been disrupted with antibiotics. In this study the injection of a mixture of essential amino acids into the haemocoel of aposymbiotic aphids was shown to alleviate, at least partially, the impact of symbiont loss. Specifically, the total amino acid content in the tissues of aposymbiotic aphids was reduced by approximately 40% to levels comparable with symbiotic insects, and there was a 1.7-fold increase in the number of embryos, suggesting that the availability of essential amino acids promotes aphid protein synthesis by rejuvenating the free amino acid pool of aposymbiotic aphids. In addition, a similar effect on the total amino acid content was observed when phenylalanine alone, but not glutamine, lysine or tryptophan, was injected into the haemocoel of aposymbiotic aphids, and there was also a significant increase in the number of embryos following injection of phenylalanine or tryptophan alone. The impact of amino acid injection on the embryo complement of aposymbiotic aphids was limited to an increase in the number of embryos, with no increase in basal embryo size. It is proposed that older embryos may rely on their own complement of symbiotic bacteria for essential amino acid provisioning. Taken together, the data highlight the importance of bacterial provisioning of essential amino acids, particularly the aromatic amino acids, in the intact symbiosis.  相似文献   

18.
Plants frequently engage in simultaneous interactions with diverse classes of biotic antagonists. Differential induction of plant defence pathways by these antagonists, and interactions between pathways, can have important ecological implications; however, these effects are currently not well understood. We explored how Tomato yellow leaf curl virus (TYLCV) influenced the performance of its vector (Bemisia tabaci) and a non‐vector herbivore (Tetranychus urticae) occurring separately or together on tomato plants (Solanum lycopersicum). TYLCV enhanced the performance of B. tabaci, although this effect was statistically significant only in the absence of T. urticae, which adversely affected B. tabaci performance regardless of infection status. In contrast, the performance of T. urticae was enhanced (only) by the combined presence of TYLCV and B. tabaci. Analyses of phytohormone levels and defence gene expression in wild‐type tomatoes and various plant‐defence mutants indicate that the enhancement of herbivore performance (for each species) entails the disruption of downstream defences in the jasmonic acid (JA) pathway. For T. urticae, this disruption appears to involve antagonistic effects of salicylic acid (SA), which is cumulatively induced to high levels by B. tabaci and TYLCV. In contrast, TYLCV was found to suppress JA‐mediated responses to B. tabaci via mechanisms independent of SA.  相似文献   

19.
The possibility that the herbicide glyphosate (N-phosphonomethylglycine) may be taken up in plant cells via a phosphate transporter of the plasma membrane was investigated using protoplasts of broad bean leaves ( Vicia faba L.). Phosphonoformic acid, a powerful inhibitor of phosphate transport in animal cells, was first demonstrated to be a competitive inhibitor of phosphate uptake inbroad bean protoplasts. Glyphosate was able to inhibit phosphate uptake into the protoplasts, and to protect partially the phosphate transporter from inhibition by phosphonoformic acid. Concentration dependence studies showed that glyphosate uptake exhibited a saturable phase at low glyphosate concentrations (0. 5 to 3 μ M ), superimposed by a linear uptake at higher concentrations (up to 100 μ M ). Inhibition of glyphosate uptake by para -chloromercuribenzene sulphonic acid, sodium azide and carbonyl-cyanide- m -chlorophenylhydrazone was much stronger at 1 than at 100 μ M glyphosate. Kinetics indicated that the saturable component of glyphosate transport was competitively inhibited by either phosphate or phosphonoformic acid. It is concluded that glyphosate can be absorbed via a phosphate transporter of the plasma membrane  相似文献   

20.
Climate adaptation has major consequences in the evolution and ecology of all living organisms. Though phytophagous insects are an important component of Earth's biodiversity, there are few studies investigating the evolution of their climatic preferences. This lack of research is probably because their evolutionary ecology is thought to be primarily driven by their interactions with their host plants. Here, we use a robust phylogenetic framework and species‐level distribution data for the conifer‐feeding aphid genus Cinara to investigate the role of climatic adaptation in the diversity and distribution patterns of these host‐specialized insects. Insect climate niches were reconstructed at a macroevolutionary scale, highlighting that climate niche tolerance is evolutionarily labile, with closely related species exhibiting strong climatic disparities. This result may suggest repeated climate niche differentiation during the evolutionary diversification of Cinara. Alternatively, it may merely reflect the use of host plants that occur in disparate climatic zones, and thus, in reality the aphid species' fundamental climate niches may actually be similar but broad. Comparisons of the aphids' current climate niches with those of their hosts show that most Cinara species occupy the full range of the climatic tolerance exhibited by their set of host plants, corroborating the hypothesis that the observed disparity in Cinara species' climate niches can simply mirror that of their hosts. However, 29% of the studied species only occupy a subset of their hosts' climatic zone, suggesting that some aphid species do indeed have their own climatic limitations. Our results suggest that in host‐specialized phytophagous insects, host associations cannot always adequately describe insect niches and abiotic factors must be taken into account.  相似文献   

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