Effect of parasitoids,seed‐predators and ant‐mutualists on fruiting success and germination of Acacia drepanolobium in Kenya

Acacia drepanolobium is an obligate ant‐plant that bears dehiscent pods exploited by predispersal seed‐predators and parasitoids. Fruit set and seed germination in relation to ant‐association, bruchid and parasitoid infestation were investigated in a large‐scale multi‐year study in Kenya. Ant‐association had a significant impact on the overall numbers of fruiting trees. 94.8% of Crematogaster mimosae‐occupied trees, 25.6% of C. nigriceps‐occupied trees, and 82.2% of Tetraponera penzigi‐occupied trees set fruit. Within each tree, ant‐association had no significant impact on the amount of seed produced: C. mimosae‐occupied trees produced 47.5 seeds per branch, C. nigriceps‐occupied trees 44.9 seeds per branch and T. penzigi‐occupied trees 38.3 seeds per branch. Ant‐association did not limit seed‐predators or vary significantly by ant. Seeds infested by bruchids germinated in significantly lower proportions (6.2%) compared to uninfested seed (78.6%). Bruchid‐infested seed is also exploited by parasitoid wasps. Parasitoids appear to have a moderate but significant ‘rescue’ effect on bruchid‐infested seed with 18.4% of parasitoid‐infested seed germinating. Stable isotopes (δ15N) revealed the trophic structure of the seed‐associated insects, showing clearly that bruchids are seed predators and the parasitoid wasp Dinarmus magnus exploits the dominant Bruchidius sp.     

Aphid infestation promotes survival of a seed predator: observations and experiments on a tritrophic community module

We investigated whether aphid presence and abundance influence the survival of an endophagous pre-dispersal seed predator of the same host plant. We studied a terrestrial community module consisting of one plant (Laburnum anagyroides) and four insect species/groups (an aphid, Aphis cytisorum, a pre-dispersal seed predator bruchid, Bruchidius villosus, aphid-attending ant species, and parasitoids of the bruchid). Two complementary investigations were carried out in parallel: (a) a plant-aphid-ant complex was experimentally manipulated by excluding aphids, ants, or both for 5 years to assess their impacts on the seed predator’s survival and parasitism rate; and (b) different aphid infestation levels on randomly selected infructescences were correlated with plant traits, nutrient allocation pattern, and variables of seed predator’s survival, such as the number of eggs laid and adults emerged influenced by parasitoid activity, for 7 years. We found that ants did not affect bruchid oviposition negatively, but egg-parasitism was significantly decreased by their presence. Plant traits, such as the number of seeds and seed mass, as well as seed predator performance were negatively affected by heavy aphid infestation. Seed predator -infested seeds had no effect on the mass of remaining seeds in the pods. This study suggests that aphids were nevertheless promoting bruchid abundance and survival, depending on their infestation rate.     

Egg dispersion is more important than competition type for herbivores attacked by a parasitoid

Herbivore fitness can be altered by a combination of interacting organisms, such as its food plant, conspecifics, and predators/parasitoids. Here, we tested relative effects of plant species, herbivore intraspecific competition type, and spatial distribution of the herbivore among plant units on herbivore survival and whether parasitoids modified these effects. We used an endophagous bruchine seed predator Callosobruchus maculatus for the herbivore, and a braconid wasp Heterospilus prosopidis for the parasitoid. The survival rate of C. maculatus was measured for each of 16 combinations of two plants (bean species, Vigna unguiculata and V. radiata), two competition types of C. maculatus larvae (contest and scramble), two spatial distributions of hosts [sparse (1 C. maculatus larva per seed over 20 seeds) and dense (2 C. maculatus larvae per seed over ten seeds)], and with/without a parasitoid pair. In the absence of the parasitoid, C. maculatus survival rate was lower with V. radiata and in the contest type. With the parasitoid, the proportion parasitized hosts was independent of total host density. Neither the proportion of parasitized hosts nor host survival rate was affected by plant species or host strain, but they were affected by host spatial distribution. When host distribution was dense, a higher proportion of hosts were parasitized, and C. maculatus survival rate was lower. Here we discuss parasitoid potential as a selective agent for the sparse within-pod distribution of its hosts in the field.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

Describing a multitrophic plant-herbivore-parasitoid system at four spatial scales

Herbivore-parasitoid interactions must be studied using a multitrophic and multispecies approach. The strength and direction of multiple effects through trophic levels may change across spatial scales. In this work, we use the herbaceous plant Ruellia nudiflora, its moth herbivore Tripudia quadrifera, and several parasitoid morphospecies that feed on the herbivore to answer the following questions: Do herbivore and parasitoid attack levels vary depending on the spatial scale considered? With which plant characteristics are the parasitoid and the herbivore associated? Do parasitoid morphospecies vary in the magnitude of their positive indirect effect on plant reproduction? We evaluated three approximations of herbivore and parasitoid abundance (raw numbers, ratios, and attack rates) at four spatial scales: regional (three different regions which differ in terms of abiotic and biotic characteristics); population (i.e. four populations within each region); patch (four 1 m² plots in each population); and plant level (using a number of plant characteristics). Finally, we determined whether parasitoids have a positive indirect effect on plant reproductive success (seed number). Herbivore and parasitoid numbers differed at three of the spatial scales considered. However, herbivore/fruit ratio and attack rates did not differ at the population level. Parasitoid/host ratio and attack rates did not differ at any scale, although there was a tendency of a higher attack in one region. At the plant level, herbivore and parasitoid abundances were related to different plant traits, varying the importance and the direction (positive or negative) of those traits. In addition, only one parasitoid species (Bracon sp.) had a positive effect on plant fitness saving up to 20% of the seeds in a fruit. These results underline the importance of knowing the scales that are relevant to organisms at different trophic levels and distinguish between the specific effects of species.     

Linkage disequilibrium at the APA insecticidal seed protein locus of common bean (<Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> L.)

Background  

An interesting seed protein family with a role in preventing insect herbivory is the multi-gene, APA family encoding the α-amylase inhibitor, phytohemagglutinin and arcelin proteins of common bean (Phaseolus vulgaris). Variability for this gene family exists and has been exploited to breed for insect resistance. For example, the arcelin locus has been successfully transferred from wild to cultivated common bean genotypes to provide resistance against the bruchid species Zabrotes subfasciatus although the process has been hampered by a lack of genetic tools for and understanding about the locus. In this study, we analyzed linkage disequilibrium (LD) between microsatellite markers at the APA locus and bruchid resistance in a germplasm survey of 105 resistant and susceptible genotypes and compared this with LD in other parts of the genome.     

Mapping of quantitative trait loci for a new source of resistance to bruchids in the wild species Vigna nepalensis Tateishi & Maxted (Vigna subgenus Ceratotropis)

Azuki bean breeders have long been interested in producing azuki bean [Vigna angularis (Willd.) Ohwi & Ohashi] varieties with bruchid resistance. A new bruchid (Callosobruchus spp.) resistance source was found in V. nepalensis Tateishi & Maxted, a species that is cross compatible with azuki bean. Quantitative trait loci (QTLs) analysis for resistance to C. chinensis (L.) and C. maculatus (F.) was conducted using F(2) (V. nepalensis x V. angularis) and BC(1)F(1) [(V. nepalensis x V. angularis) x V. angularis] populations derived from crosses between the bruchid resistant species V. nepalensis and bruchid susceptible species V. angularis. Resistance was measured using two traits, percentage of seeds damaged by bruchids and the time taken for adult bruchids to emerge from seeds. Based on the results from both populations seven QTLs were detected for bruchid resistance; five QTLs for resistance to C. chinensis and two QTLs for resistance to C. maculatus. The different locations found for some resistance QTL to the two bruchid species suggests different resistance mechanisms. QTLs on linkage group (LG) 1 and LG2 for bruchid resistance to C. chinensis co-localized with seed size QTLs suggesting that incremental increase in seed size accompanied susceptibility to C. chinensis. Based on linked markers the QTL on these two linkage groups appear to be the same as previously reported in other Asian Vigna. However, several other QTLs were newly detected including one on LG4 that appears unrelated to seed size. Transfer of these new sources of bruchid resistance from V. nepalensis to azuki bean will be aided by the progress being made in azuki genome mapping.     

Interaction between ungulates and bruchid beetles and its effect on <Emphasis Type="Italic">Acacia</Emphasis> trees: modeling the costs and benefits of seed dispersal to plant demography

Integrative studies of plant–animal interactions that incorporate the multiple effects of interactions are important for discerning the importance of each factor within the population dynamics of a plant species. The low regeneration capacity of many Acacia species in arid savannas is a consequence of a combination of reduction in seed dispersal and high seed predation. Here we studied how ungulates (acting as both seed dispersers and herbivores) and bruchid beetles (post-dispersal seed predators) modulate the population dynamics of A. raddiana, a keystone species in the Middle East. We developed two simulation models of plant demography: the first included seed ingestion by ungulates and seed predation by bruchids, whereas the second model additionally incorporated herbivory by ungulates. We also included the interacting effects of seed removal and body mass, because larger ungulates destroy proportionally fewer seeds and enhance seed germination. Simulations showed that the negative effect of seed predation on acacia population size was compensated for by the positive effect of seed ingestion at 50 and 30% seed removal under scenarios with and without herbivory, respectively. Smaller ungulates (e.g., <35 kg) must necessarily remove tenfold more seeds than larger ungulates (e.g., >250 kg) to compensate for the negative effect of seed predation. Seedling proportion increased with seed removal in the model with herbivory. Managing and restoring acacia seed dispersers is key to conserving acacia populations, because low-to-medium seed removal could quickly restore their regeneration capacity.     

The interaction between host and host plant influences the oviposition and performance of a generalist ectoparasitoid

The successful development of parasitoids of herbivores depends on the quality of their host, which is often affected by the host plant. Therefore, a parasitoid’s oviposition decisions will directly depend on the host, but also on plant quality. Here, we investigated the direct effects of host species and the indirect effects of the host’s food plant on the oviposition decisions and performance of the gregarious ectoparasitoid Euplectrus platyhypenae Howard (Hymenoptera: Eulophidae). With a series of no‐choice experiments, we determined the oviposition and performance of the parasitoid on: (1) two caterpillar species, fall armyworm, Spodoptera frugiperda JE Smith (Lepidoptera: Noctuidae), and velvet armyworm, Spodoptera latifascia Walker, reared on maize (Zea mays L., Poaceae), (2) the same caterpillars reared on maize, bean (Phaseolus vulgaris L., Fabaceae), or squash (Cucurbita pepo L., Cucurbitaceae) leaves, and (3) S. latifascia caterpillars reared on leaves of wild and cultivated lima bean, Phaseolus lunatus L. All these insects and plants originate from Mesoamerica where they have coexisted for thousands of years in the traditional agricultural system known as Milpa in which maize, beans, and squash are planted together. We found that the preferred and best combination of host and host plant for parasitoid performance was S. frugiperda on maize. Parasitoids laid larger clutches, had higher survival, and more females and larger adults emerged from S. frugiperda reared on maize. However, when both caterpillar species were reared on squash, S. latifascia was the better host. Contrary to the literature, S. frugiperda was not able to develop on bean plants. Results from the lima bean experiment showed that parasitoid performance was best when S. latifascia was reared on leaves of cultivated compared to wild lima bean. These findings are discussed in the context of mixed cropping in which the ability of generalist parasitoids to switch among hosts and host plant species could be advantageous for pest management.     

Spatial Variation in the Strength of a Trophic Cascade Involving Ruellia nudiflora (Acanthaceae), an Insect Seed Predator and Associated Parasitoid Fauna in Mexico

Spatial variation in the strength of herbivore top-down control represents an important source of variation in plant fitness measures and community structure and function. By measuring seed predator (larvae of a Noctuid moth) and parasitoid impacts on Ruellia nudiflora across a broad spatial scale in Yucatan (Mexico), this study addressed the following: (1) to what extent does seed predator and parasitoid attack intensity associated with R. nudiflora vary spatially? (2) Does parasitoid attack result in a positive indirect effect on the plant, and does the intensity of this effect vary spatially? During the peak of fruit production (late June–early July) of 2005, we collected fruits from 21 R. nudiflora populations and grouped them into four regions: center, east, north and south. For each fruit we recorded: observed seed number, number of seeds eaten, seed predator presence, parasitoid presence and number of seeds ‘saved’ by parasitoids. Seed predators attacked ca 30 percent of fruits/plant on average, while parasitoids were found in 24 percent of seed predator-attacked fruits. Results indicated spatial variation in seed predator and parasitoid attack levels; interestingly, a contrasting spatial gradient of attack intensity was observed: populations/regions with greatest parasitoid attack levels usually had the lowest seed predator attack levels and vice versa, suggesting top-down control of parasitoids on seed predators. We observed a weak overall indirect impact of parasitoids on R. nudiflora (4% seeds ‘saved’ on average), which nonetheless varied strongly across populations (e.g., close to 14% seeds saved at one population). Findings indicate a geographical structuring of interaction strengths across populations, as well as spatial variation in the strength of parasitoid cascading effects on plant reproduction.     

Characterization of digestive enzymes of bruchid parasitoids-initial steps for environmental risk assessment of genetically modified legumes

Genetically modified (GM) legumes expressing the α-amylase inhibitor 1 (αAI-1) from Phaseolus vulgaris L. or cysteine protease inhibitors are resistant to several bruchid pests (Coleoptera: Chrysomelidae). In addition, the combination of plant resistance factors together with hymenopteran parasitoids can substantially increase the bruchid control provided by the resistance alone. If the strategy of combining a bruchid-resistant GM legume and biological control is to be effective, the insecticidal trait must not adversely affect bruchid antagonists. The environmental risk assessment of such GM legumes includes the characterization of the targeted enzymes in the beneficial species and the assessment of the in vitro susceptibility to the resistance factor. The digestive physiology of bruchid parasitoids remain relatively unknown, and their susceptibility to αAI-1 has never been investigated. We have detected α-amylase and serine protease activities in all five bruchid parasitoid species tested. Thus, the deployment of GM legumes expressing cysteine protease inhibitors to control bruchids should be compatible with the use of parasitoids. In vitro inhibition studies showed that sensitivity of α-amylase activity to αAI-1 in the parasitoids was comparable to that in the target species. Direct feeding assays revealed that harmful effects of α-amylase inhibitors on bruchid parasitoids cannot be discounted and need further evaluation.     

Embryo abortion and histological features in the interspecific cross between <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> L. and <Emphasis Type="Italic">P. coccineus</Emphasis> L

In this paper, the causes of early embryo abortion in the reciprocal crosses between Phaseolus vulgaris L. (a cultivar) and Phaseolus coccineus L. (a wild form) were studied. Methacrylate resin sections, 3–5 μm thick, of 3 to 14 day-old seeds were used to examine the embryo developmental stages and the state of seed tissue. It was observed that, embryos aborted at different developmental stages (globular to early cotyledon) depending on the maternal parent. The use of P. coccineus cytoplasm resulted in a higher number of abortion than in reciprocal crosses. Many of them took place between 5 and 6 days after pollination (DAP). Histological analyses permitted to observe that the embryo development was slower in the cross between P. coccineus and P. vulgaris, compared to parental seeds. It would be related to a deficient endosperm development in reciprocal crosses and, in some extent, hypertrophy of the suspensor might be the main cause of early embryo abortion. Then, it would be practical to overcome this incompatibility by rescuing the embryo at the globular stage of development.     

Extended larval development time for aphid parasitoids in the presence of plant endosymbionts

Abstract 1. Variation in plant chemistry does not only mediate interactions between plants and herbivores but also those between herbivores and their natural enemies, and plants and natural enemies. 2. Endophytic fungi complete their whole life cycle within the host plant’s tissue and are associated with a large diversity of plant species. Endophytes of the genus Neotyphodium alter the chemistry of the host plant by producing herbivore toxic alkaloids. 3. Here we asked whether the endophyte‐tolerant aphid species Metopolophium festucae could be defended against its parasitoid Aphidius ervi when feeding on endophyte‐infected plants. In a laboratory experiment, we compared life‐history traits of A. ervi when exposed to hosts on endophyte‐infected or endophyte‐free Lolium perenne. 4. The presence of endophytes significantly increased larval and pupal development times, but did not affect the mortality of immature parasitoids or the longevity of the adults. Although the number of parasitoid mummies tended to be reduced on endophyte‐infected plants, the number of emerging parasitoids did not differ significantly between the two treatments. 5. This shows that the metabolism of individual aphids feeding on infected plants may be changed and help in the defence against parasitoids. An increase in parasitoid development time should ultimately reduce the population growth of A. ervi. Therefore, endophyte presence may represent an advantage for endophyte‐tolerant aphid species through extended parasitoid development and its effect on parasitoid population dynamics.     

Differential induction of plant chemical defenses by parasitized and unparasitized herbivores: consequences for reciprocal,multitrophic interactions

Insect parasitoids can play ecologically important roles in virtually all terrestrial plant–insect herbivore interactions, yet whether parasitoids alter the defensive traits that underlie interactions between plants and their herbivores remains a largely unexplored question. Here, we examined the reciprocal trophic interactions among populations of the wild cabbage Brassica oleracea that vary greatly in their production of defensive secondary compounds – glucosinolates (GSs), a generalist herbivore, Trichoplusia ni, and its polyembryonic parasitoid Copidosoma floridanum. In a greenhouse environment, plants were exposed to either healthy (unparasitized), parasitized, or no herbivores. Feeding damage by herbivores induced higher levels of the indole GSs, glucobrassicin and neoglucobrassicin, but not any of the other measured GSs. Herbivores parasitized by C. floridanum induced cabbage plants to produce 1.5 times more indole GSs than levels induced by healthy T. ni and five times more than uninduced plants. As a gregarious endoparasitoid, C. floridanum causes its host T. ni to feed more than unparasitized herbivores resulting in increased induction of indole GSs. In turn, herbivore fitness parameters (including differential effects on male and female contributions to lifetime fecundity in the herbivore) were negatively correlated with the aliphatic GSs, sinigrin and gluconapin, whereas parasitoid fitness parameters were negatively correlated with the indole GSs, glucobrassicin and neoglucobrassicin. That herbivores and their parasitoids appear to be affected by different sets of GSs was unexpected given the intimate developmental associations between host and parasitoid. This study is the first to demonstrate that parasitoids, through increasing feeding by their herbivorous hosts, can induce higher levels of non‐volatile plant chemical defenses. While parasitoids are widely recognized to be ubiquitous in most terrestrial insect herbivore communities, their role in influencing plant–insect herbivore relationships is still vastly underappreciated.     

Impact of αAI-1 Expressed in Genetically Modified Cowpea on Zabrotes subfasciatus (Coleoptera: Chrysomelidae) and Its Parasitoid,Dinarmus basalis (Hymenoptera: Pteromalidae)

Genetically modified (GM) cowpea seeds expressing αAI-1, an α-amylase inhibitor from the common bean, have been shown to be immune against several bruchid species. Effective control of such pests by growing GM cowpea could promote the spread of bruchid species that are αAI-1 tolerant. Consequently, the sustainability of bruchid pest control could be increased by combining GM seeds and hymenopteran parasitoids. However, there are concerns that αAI-1 could interfere with the biological control provided by parasitoids. Here, we assessed the impact of GM cowpea seeds expressing αAI-1 on the αAI-1-tolerant bruchid Zabrotes subfasciatus and its parasitoid Dinarmus basalis. αAI-1 in cowpea seeds did not increase resistance to Z. subfasciatus or affect the mortality rate of Z. subfasciatus larvae. Parasitism of Z. subfasciatus by D. basalis and fitness of D. basalis offspring were not affected by the presence of αAI-1. Thus, αAI-1-expressing cowpeas and parasitoids should be compatible for the control of bruchid pests.     

Intraspecific variation in herbivore‐induced plant volatiles influences the spatial range of plant–parasitoid interactions

Chemical information influences the behaviour of many animals, thus affecting species interactions. Many animals forage for resources that are heterogeneously distributed in space and time, and have evolved foraging behaviour that utilizes information related to these resources. Herbivore‐induced plant volatiles (HIPVs), emitted by plants upon herbivore attack, provide information on herbivory to various animal species, including parasitoids. Little is known about the spatial scale at which plants attract parasitoids via HIPVs under field conditions and how intraspecific variation in HIPV emission affects this spatial scale. Here, we investigated the spatial scale of parasitoid attraction to two cabbage accessions that differ in relative preference of the parasitoid Cotesia glomerata when plants were damaged by Pieris brassicae caterpillars. Parasitoids were released in a field experiment with plants at distances of up to 60 m from the release site using intervals between plants of 10 or 20 m to assess parasitism rates over time and distance. Additionally, we observed host‐location behaviour of parasitoids in detail in a semi‐field tent experiment with plant spacing up to 8 m. Plant accession strongly affected successful host location in field set‐ups with 10 or 20 m intervals between plants. In the semi‐field set‐up, plant finding success by parasitoids decreased with increasing plant spacing, differed between plant accessions, and was higher for host‐infested plants than for uninfested plants. We demonstrate that parasitoids can be attracted to herbivore‐infested plants over large distances (10 m or 20 m) in the field, and that stronger plant attractiveness via HIPVs increases this distance (up to at least 20 m). Our study indicates that variation in plant traits can affect attraction distance, movement patterns of parasitoids, and ultimately spatial patterns of plant–insect interactions. It is therefore important to consider plant‐trait variation in HIPVs when studying animal foraging behaviour and multi‐trophic interactions in a spatial context.     

Food plant and herbivore host species affect the outcome of intrinsic competition among parasitoid larvae

1. In nature, several parasitoid species often exploit the same stages of a common herbivore host species and are able to coexist despite competitive interactions amongst them. Less is known about the direct effects of resource quality on intrinsic interactions between immature parasitoid stages. The present study is based on the hypothesis that variation in the quality or type of plant resources on which the parasitoids indirectly develop may be complementary and thus facilitate niche segregation favouring different parasitoids in intrinsic competition under different dietary regimes. 2. The present study investigated whether two herbivore species, the cabbage butterflies Pieris brassicae and Pieris rapae (Pieridae), and the quality of two important food plants, Brassica oleracea and Brassica nigra (Brassicaceae), affect the outcome of intrinsic competition between their primary larval endoparasitoids, the gregarious Cotesia glomerata (Braconidae) and the solitary Hyposoter ebeninus (Ichneumonidae). 3. Hyposoter ebeninus is generally an intrinsically superior competitor over C. glomerata. However, C. glomerata survived more antagonistic encounters with H. ebeninus when both developed in P. brassicae rather than in P. rapae caterpillars, and while its host was feeding on B. nigra rather than B. oleracea. Moreover, H. ebeninus benefitted from competition by its higher survival in multiparasitised hosts. 4. These results show that both plant and herbivore species mediate the battleground on which competitive interactions between parasitoids are played out and may affect the outcomes of these interactions in ways that enable parasitoids to segregate their niches. This in turn may promote coexistence among parasitoid species that are associated with the same herbivore host.     

Mycorrhiza modulates aboveground tri‐trophic interactions to the fitness benefit of its host plant

1. Arbuscular mycorrhiza (AM), the association of AM fungi and plant roots, may alter morphological and physiological attributes of aboveground plant parts and thereby influence plant‐associated organisms such as herbivores and their natural enemies, predators and parasitoids. 2. The interactions between AM and the players of aboveground tri‐trophic systems have mainly been considered in isolation from each other. The effects of AM on aboveground herbivore–carnivore population dynamics and the consequences to plant fitness are unknown. 3. We explored AM‐induced compensatory mechanisms for AM‐promoted proliferation of the herbivorous spider mite, Tetranychus urticae Koch, on whole bean plants, Phaseolus vulgaris L. Vegetative and reproductive plant growth, AM fungal colonisation levels, and mite densities were assessed on spider mite‐infested plants colonised or not by the AM fungus Glomus mosseae Nicol. & Gerd, and harbouring the natural enemy of the spider mites, the predatory mite Phytoseiulus persimilis Anthias‐Henriot or not. 4. AM symbiosis modulated the aboveground tri‐trophic system to the fitness benefit of the plant. AM‐increased plant productivity outweighed the fitness decrease due to AM‐promoted herbivory: at similar vegetative growth, mycorrhizal plants produced more seeds than non‐mycorrhizal plants. 5. AM‐increased spider mite population levels were compensated for by enhanced population growth of the predators and increased plant tolerance to herbivory. 6. AM‐enhanced predator performance looped back to the AM fungus and stabilised its root colonisation levels, providing the first experimental evidence of a mutually beneficial interaction between AM and an aboveground third trophic level natural enemy.     

Altering protein composition by genetically removing phaseolin from common bean seeds containing arcelin or phytohemagglutinin

 Arcelin seed proteins of common bean (Phaseolus vulgaris L.) confer resistance to bruchid pests, and in vitro results suggested that greater resistance could be achieved by increasing the concentration of arcelin. We created backcross lines having arcelin alleles (SMARC lines), or alleles for the related protein phytohemagglutinin (PHA) (SMPHA lines), and a null allele for phaseolin to determine if seeds lacking phaseolin would contain increased quantities of arcelin or PHA proteins. To test the affects of genetically removing phaseolin, SMARC and SMPHA lines were derived as pairs of phaseolin-containing and phaseolin-null lines. Parental, SMARC, and SMPHA lines were grown in a replicated greenhouse trial and measured for days-to-flower, days-to-maturity, seed weight, and for quantities of phaseolin, arcelin dimer, PHA, and total proteins. There were no differences between pairs of phaseolin and phaseolin-null lines for days-to-flower, seed weight or total protein, and inconsistent differences for days-to-maturity. Arcelin concentrations were significantly increased in two of four pairs of SMARC lines, and PHA concentration was significantly greater in four of five pairs of SMPHA lines. These or other changes in the seed protein composition in phaseolin null lines may improve resistance to bruchids. Received: 2 April 1997 / Accepted: 20 May 1997     

Effects of plant sex on insect abundance across three trophic levels in the perennial shrub Buddleja cordata

Although there is a growing interest in the effects of intra‐specific plant genetic variation on species interactions, the effects of plant sex, an important axis of genetic variation, have been less studied. In addition, previous work investigating plant sex effects on species interactions has frequently focused on bitrophic interactions (e.g., herbivory), usually ignoring plant sex effects on higher trophic levels (i.e., natural enemies). Here, we investigated the effects of plant sex on herbivore abundance and that of their natural enemies associated with the dioecious shrub Buddleja cordata Kunth (Scrophulariaceae). Furthermore, we measured a subset of plant traits frequently involved in herbivore resistance and the potentially underlying plant sex effects. To this end, we recorded the abundances of a specialist leaf‐chewing caterpillar [Acronyctodes mexicanaria Walker (Lepidoptera: Geometridae)] throughout an entire growing season. We also recorded information about the caterpillar’s parasitoids, as well as leaf water content, phenolic compounds, phosphorus, and nitrogen for male and female plants of B. cordata. Plant sex did not significantly influence caterpillar abundance but did have an effect on natural enemies, with parasitoid abundance being 2.4‐fold greater on female than on male plants. The effect of plant sex on parasitoids remained significant after accounting for caterpillar abundance, suggesting that it was underlain by a trait‐mediated (rather than density‐mediated) mechanism. Finally, we found that male plants had a higher concentration of phenolic compounds (other traits did not differ between plant sexes). These results provide valuable evidence for the extended effects of plant sex on the third trophic level and point at plant traits potentially mediating such effects.