Effects of population-related variation in plant primary and secondary metabolites on aboveground and belowground multitrophic interactions

Insects feeding on aboveground and belowground tissues can influence each other through their shared plant and this is often mediated by changes in plant chemistry. We examined the effects of belowground root fly (Delia radicum) herbivory on the performance of an aboveground herbivore (Plutella xylostella) and its endoparasitoid wasp (Cotesia vestalis). Insects were reared on three populations of wild cabbage (Brassica oleracea) plants, exhibiting qualitative and quantitative differences in root and shoot defense chemistry, that had or had not been exposed to root herbivory. In addition, we measured primary (amino acids and sugars) and secondary [glucosinolate (GS)] chemistry in plants exposed to the various plant population-treatment combinations to determine to what extent plant chemistry could explain variation in insect performance variables using multivariate statistics. In general, insect performance was more strongly affected by plant population than by herbivory in the opposite compartment, suggesting that population-related differences in plant quality are larger than those induced by herbivory. Sugar profiles were similar in the three populations and concentrations only changed in damaged tissues. In addition to population-related differences, amino acid concentrations primarily changed locally in response to herbivory. Whether GS concentrations changed in response to herbivory (indole GS) or whether there were only population-related differences (aliphatic GS) depended on GS class. Poor correlations between performance and chemical attributes made biological interpretation of these results difficult. Moreover, trade-offs between life history traits suggest that factors other than food nutritional quality contribute to the expression of life history traits.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

Ecological implications of anti-pathogen effects of tropical fungal endophytes and mycorrhizae

We discuss studies of foliar endophytic fungi (FEF) and arbuscular mycorrhizal fungi (AMF) associated with Theobroma cacao in Panama. Direct, experimentally controlled comparisons of endophyte free (E-) and endophyte containing (E+) plant tissues in T. cacao show that foliar endophytes (FEF) that commonly occur in healthy host leaves enhance host defenses against foliar damage due to the pathogen (Phytophthora palmivora). Similarly, root inoculations with commonly occurring AMF also reduce foliar damage due to the same pathogen. These results suggest that endophytic fungi can play a potentially important mutualistic role by augmenting host defensive responses against pathogens. There are two broad classes of potential mechanisms by which endophytes could contribute to host protection: (1) inducing or increasing the expression of intrinsic host defense mechanisms and (2) providing additional sources of defense, extrinsic to those of the host (e.g., endophyte-based chemical antibiosis). The degree to which either of these mechanisms predominates holds distinct consequences for the evolutionary ecology of host-endophyte-pathogen relationships. More generally, the growing recognition that plants are composed of a mosaic of plant and fungal tissues holds a series of implications for the study of plant defense, physiology, and genetics.     

Jasmonic acid treatment and mammalian herbivory differentially affect chemical defenses and growth of wild mustard (Brassica kaber)

Summary. Jasmonic acid (JA) is a wound-related hormone found in most plants that, when applied exogenously, can induce increases in levels of chemical defenses in patterns similar to those induced by mechanical damage or insect feeding. Relative to responses to insect and pathogen attack, chemical responses of herbaceous plants to mammalian herbivore attack have been little studied. In a field experiment, we compared the effects of JA treatment and naturally occurring mammalian herbivory on the expression of trypsin inhibitors, glucosinolates, peroxidase activity and growth of wild mustard (Brassica kaber). Exogenous JA significantly increased trypsin inhibitor activity and glucosinolate concentration, and moderately increased peroxidase activity in the eighth true leaves of five-week-old plants, relative to untreated controls. In contrast, levels of these chemical defenses in the eighth true leaves or in regrowth foliage of plants that had ∼80% of their leaf area removed by groundhogs (Marmota monax) did not differ from that in undamaged and untreated controls. Although exogenous JA significantly elevated levels of chemical defenses, it did not affect height of plants through the season and only slightly reduced time to first flower. Groundhog herbivory significantly reduced height and delayed or abolished flowering, but these effects were not substantial unless coupled with apical meristem removal. We hypothesize that the lack of effect of groundhog herbivory on chemical defenses may be due in part to the speed and pattern of leaf area removal by groundhogs, or physiological constraints caused by leaf area loss. Despite having no effect on chemical defense production, leaf area loss by groundhogs was more costly to growth and fitness than the effects of JA application in this study, but only substantially so if coupled with apical meristem removal. We suggest that in general, costs of defense production in plants are likely to be minimal when compared to the risk of losing large amounts of leaf area or primary meristematic tissue. Thus, if they are effective at deterring herbivory, the benefits of inducible defense production likely outweigh the costs in most cases. Received 20 December 2000; accepted 3 May 2001     

两种光照生境下4种常绿阔叶树的单位叶面积干重、光合能力与化学防御物质含量比较

亚热带常绿阔叶林下分布着可塑性强的耐荫植物,其叶片应对外界干扰的防御对策至少有物理性防御和化学性防御两种,这两种防御代价在一定的昆虫取食压力下在理论上应该存在权衡.为检验这种权衡是否普遍存在,分别测定了重庆缙云山4个常绿物种——光叶山矾(Symplocos lancifolia)、四川山矾(S.setchuensis)、四川毛蕊茶(Camellialawii)和细枝柃(Eurya loquaiana)在林窗和林下两种对照光环境下个体叶片的光饱和下的净光合速率、叶面积干重和部分化学防御物质含量.结果表明,4个物种生活在林窗的个体单位叶面积干重、叶片氮含量和单位面积净光合速率通常高于林下的个体,但是单位质量和单位面积的总酚含量和C/N值却低于林下的个体.蛋白质和非结构性碳水化合物含量在不同生境中没有显著差异.因为不同物种叶片对光照条件的响应不同,不同参数的变化在两种生境间的差异并不完全一致.本研究结果在一定程度上暗示了物种在林窗和林下分别倾向于采取物理和化学性防御对策,并初步证实了两种防御强度之间存在权衡关系.图4参40     

Induced responses to herbivory in the Neotropical ant-plant association between Azteca ants and Cecropia trees: response of ants to potential inducing cues

Plant defense against herbivores often involves constitutive and inducible mechanisms of resistance. Obligate ant-plants, which provide food and housing for ants, are thought to primarily rely on ants for defense against herbivores. This form of plant defense has largely been viewed as static. We have been investigating the dynamic nature of Azteca ants as an inducible defense of Cecropia trees. Ants rapidly recruit to and patrol sites of foliar damage. We propose that Azteca ants can be viewed as an inducible defense for Cecropia trees because of their sensitivity to cues associated with herbivory, their rapid and aggressive recruiting ability, and their reclaimable and redeployable nature as a plant defense. In this study, we examine ant behavior following plant damage, and the potential cues that indude ant recruitment. We found that ants present on leaves when the plant is damaged leave the damaged leaf and recruit other ants to it, presumably by laying recruitment trails. Volatile leaf cues associated with herbivory were important in eliciting an induced response in two experiments. However, we found that cues associated with a congeneric plant elicited a much stronger ant response than conspecific cues. Although the type of leaf damage (gaping wounds versus leaf edge wounds) did not affect the level of ant recruitment, the extent of damage did. Leaves with one hole punched showed a 50% increase in ants, while leaves with five holes punched in them elicited a 100% increase in ant numbers. In sum, it appears that multiple plant-related cues associated with herbivory are involved in induction of ant recruitment in the Cecropia-Azteca system. We discuss the generality of ant responses to herbivory in obligate ant-plant systems, and in facultative ant-plant associations, which may be more common. Received: 23 March 1998 / Accepted after revision: 5 July 1998     

Allocation of herbivory-induced hydroxamic acids in the wild wheat Triticum uniaristatum

Summary. We characterized the induction of hydroxamic acids (Hx) by aphid infestation in the wild wheat Triticum uniaristatum by addressing the following questions: i) Do different leaves have similar responses to aphid damage?, ii) Is the Hx induction localized or systemic?, iii) How long does the induction last?, and iv) Is the degree of damage related to the magnitude of induced Hx? Based on earlier results on this wheat/aphid system (lack of costs of Hx induction) we expected to find the plant exhibiting cost-saving patterns of response to herbivory. Aphid infestation in the primary leaf led to induced levels of Hx, but no differences in Hx levels were found after infestation of the secondary leaf. Induction of Hx was restricted to the infested leaf (primary leaf). Induced Hx levels exhibited by the primary leaf at the end of aphid infestation were not observed 2 days later. Finally, different aphid densities (between 10 and 40 aphids per leaf) did not produce significant differences in Hx levels in infested primary leaves. Characteristics of Hx induction by aphid infestation in T. uniaristatum partially support the expected cost-saving patterns in the allocation of induced defenses. Received 15 January 1997; accepted 7 July 1997.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Attraction of entomopathogenic nematodes to sugarcane root volatiles under herbivory by a sap-sucking insect

Few systems have been described in which herbivore-induced root volatiles mediate attraction of entomopathogenic nematodes (EPNs), and they only concern root damage inflicted by chewing insects. EPNs, especially Heterorhabditis indica and Steinernema carpocapsae, are potential biological control agents of sugarcane spittlebug (Mahanarva fimbriolata) populations. Here, we investigated the response of these two species of EPNs to sugarcane root volatiles damaged by M. fimbriolata nymphs in a belowground six-arm olfactometer. We also examined changes on root volatile profile in response to herbivory of sugarcane spittlebug nymphs. Results showed that both EPN species did not discriminate between odors of undamaged sugarcane and moistened sand (blank). However, when EPNs were exposed to odors of spittlebug-damaged and undamaged sugarcane roots, both species significantly preferred odors of spittlebug-damaged roots. Headspace collection followed by GC–MS analyses showed no qualitative difference (total of 11 compounds) between volatile profiles of spittlebug-damaged and undamaged sugarcane roots. In contrast to the previous studies involving feeding by root chewing insects, our root volatile analysis did not reveal any up-regulation resulting from sugarcane spittlebug damage, but the down-regulation of the terpenes dihydromyrcenol and β-isomethyl ionone when compared with the profile of undamaged sugarcane roots. Here, we propose alternative explanations for the EPN attraction to spittlebug-damaged roots as it is unlikely that reduced concentrations of the volatiles play a role in this interaction. Further studies are necessary to determine the key compounds of the root volatile emission to enhance biological control efficacy with EPNs against M. fimbriolata in sugarcane.     

发根农杆菌T—DNA基因对3种葛属植物毛状根形态和葛根素含量的影响

野葛、山葛和三裂叶葛的离体叶片与发展根农杆菌R1601直接感染诱导出毛状根并能够合成葛根素，毛状根离体培养后均出现两种形态：一种是有较高的葛根素含量的典型毛状根，另一种是葛根素含量低但生长速度较快的愈伤组织化毛状根，两类毛状根均有rol/B基因存在，表明rolB基因对所有毛状根的诱导和生长是必需的，aux1存在于所有愈伤组织化毛状根，而在典型毛状根中的检出率为20％－50％，表明aux1基因有诱导毛状根产生愈作组织的作用，含有ags基因的毛状根葛根素含量低于不含ags基因的毛状葛根素含量，表明ags基因不利于次生代谢合成，表3参12     

Making scents of defense: do fecal shields and herbivore-caused volatiles match host plant chemical profiles?

Many plant families have aromatic species that produce volatile compounds which they release when damaged, particularly after suffering herbivory. Monarda fistulosa (Lamiaceae) makes and stores volatile essential oils in peltate glandular trichomes on leaf and floral surfaces. This study examined the larvae of a specialist tortoise beetle, Physonota unipunctata, which feed on two M. fistulosa chemotypes and incorporate host compounds into fecal shields, structures related to defense. Comparisons of shield and host leaf chemistry showed differences between chemotypes and structures (leaves vs. shields). Thymol chemotype leaves and shields contained more of all compounds that differed than did carvacrol chemotypes, except for carvacrol. Shields had lower levels of most of the more volatile chemicals than leaves, but more than twice the amounts of the phenolic monoterpenes thymol and carvacrol and greater totals. Additional experiments measured the volatiles emitted from M. fistulosa in the absence and presence of P. unipunctata larvae and compared the flower and foliage chemistry of plants from these experiments. Flowers contained lower or equal amounts of most compounds and half the total amount, compared to leaves. Plants subjected to herbivory emitted higher levels of most volatiles and 12 times the total amount, versus controls with no larvae, including proportionally more of the low boiling point chemicals. Thus, chemical profiles of shields and volatile emissions are influenced by the amounts and volatilities of compounds present in the host plant. The implications of these results are explored for the chemical ecology of both the plant and the insect.     

Constitutive or induced defences - how does <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> defend itself from larval feeding?

Summary. Following herbivory, induced responses involving plant secondary metabolites have been reported in a number of tree species. Although a wide range of plant secondary metabolites appear to operate as constitutive plant defences in trees belonging to the Eucalyptus genus, no induced responses have as yet been reported following foliar-chewing insect damage. We empirically tested whether branch defoliation (artificial and larval) of 2-year-old Eucalyptus globulus Labill. trees altered the abundance of specific plant secondary metabolites immediately (3 months after initial larval feeding) and 8 months after the cessation of larval feeding. Metabolites assayed, included essential oils, polyphenolic groups and foliar wax compounds and in all cases their abundance was not significantly altered by defoliation. However, the level of foliar tannins after 3 months of larval feeding did display a trend that suggested elevated levels as the result of defoliation, though this trend was not evident 8 months later, indicating that, if real, the response was a rapid and not a delayed induced response. The level of foliar tannins was also negatively correlated to both average larval survival and average percentage branch defoliation, suggesting that foliar tannins may operate as toxins and/or anti-feedants to M. privata larval feeding.     

Indirect benefits of symbiotic coccoids for an ant-defended myrmecophytic tree

The net benefits of mutualism depend directly on the costs and effectiveness of mutualistic services and indirectly on the interactions that affect those services. We examined interactions among Cordia alliodora myrmecophytic trees, their symbiotic ants Azteca pittieri, coccoid hemipterans, and foliar herbivores in two Neotropical dry forests. The tree makes two investments in symbiotic ants: it supplies nesting space, as domatia, and it provides phloem to coccoids, which then produce honeydew that is consumed by ants. Although higher densities of coccoids should have higher direct costs for trees, we asked whether higher densities of coccoids can also have higher indirect benefits for trees by increasing the effectiveness of ant defense against foliar herbivores. We found that trees benefited from ant defense against herbivores. Ants defended trees effectively only when colonies reached high densities within trees, and ant and coccoid densities within trees were strongly positively correlated. The benefits of reduced foliar herbivory by larger ant colonies were therefore indirectly controlled by the number of coccoids. Coccoid honeydew supply also affected per capita ant aggression against tree herbivores. Ants experimentally fed a carbohydrate-rich diet, analogous to sugar obtained from coccoids, were more aggressive against caterpillars per capita than ants fed a carbohydrate-poor diet. Ant defense was more effective on more valuable and vulnerable young leaves than on older leaves. Young domatia, associated with young leaves, contained higher coccoid densities than older domatia, which suggests that coccoids may also drive spatially favorable ant defense of the tree. If higher investments by one mutualistic partner are tied to higher benefits received from the other, there may be positive feedback between partners that will stabilize the mutualism. These results suggest that higher investment by trees in coccoids leads to more effective defense by ants against the tree's foliar herbivores.     

Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

无机硅叶面肥及土壤调理剂对水稻铅、镉吸收的影响

喷施叶面阻控剂和施用土壤调理剂是目前污染稻田治理的主要措施,针对不同污染类型、污染程度稻田研发效果较好的叶面阻控剂及土壤调理剂是研究重点。文章通过2019年的田间试验研究无机硅叶面肥及土壤调理剂对中重度铅镉复合污染稻田水稻铅镉阻控效果。共设3个处理:常规施肥处理(CK)、常规施肥+喷施无机硅叶面肥处理(L-Si)、常规施肥+施用无机硅土壤调理剂处理(S-Si)。结果表明:L-Si处理土壤中铅质量分数较对照(116.00±4.35)mg·kg-1高18.97%;S-Si处理较对照(5.51±0.06)能够显著提高稻田土壤pH值0.73,土壤中铅和镉分别较对照(116.00±4.35)、(1.89±0.03)mg·kg-1高12.07%和14.81%。L-Si处理下铅在根、叶片、稻米中富集较对照分别减少11.69%、12.40%和38.46%;镉在叶片、稻壳、稻米中较对照显著减少了18.10%、5.84%和40.84%。而S-Si处理中铅在根、茎、叶、稻壳、稻米中富集量较对照显著减少11.69%、36.93%、15.16%、23.47%和51.28%;镉在根、茎、叶、稻壳、稻米中富集量显著降低12.93%、42.28%、61.75%、59.85%和49.21%。L-Si及S-Si处理水稻产量较对照(7052.27±95.07)kg·hm^-2分别提高3.24%和4.09%。综上所述,无机硅经叶面吸收后,在减少叶片中铅富集的同时减少了根系对土壤铅的吸收,将经根系吸收的铅富集于茎和稻壳中;而在减少叶片中镉富集的同时并未减少根系对镉吸收,将经根系吸收的镉富集于根和茎中。无机硅土壤调理剂在将土壤中铅和镉固定于土壤中的同时阻隔了水稻根系对铅和镉下吸收,因此水稻根、茎、叶、稻壳、稻米中铅和镉的含量均显著降低。     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

How flexible is phenotypic plasticity? Developmental windows for trait induction and reversal

Inducible defenses allow prey to modulate their phenotypic responses to the level of predation risk in the environment and reduce the cost of constitutive defenses. Inherent in this statement is that prey must alter their phenotypes during development in order to form these defenses. This has lead many ecologists and evolutionary biologists to call for studies that examine developmental plasticity to provide insights into the importance of development in controlling the trajectories of trait formation, the integration of phenotypes over ontogeny, and the establishment of developmental windows for trait formation and reversal. By moving away from studies that focus on a single point in development, we can obtain a more complete understanding of the phenotypic decisions and limitations of prey. We exposed freshwater snails (Helisoma trivolvis) to environments in which predatory water bugs (Belostoma flumineum) were always absent, always present, or added and removed at different points in development. We discovered that snails formed morphological defenses against water bugs. Importantly, after the initial induction of defenses, snails showed similar developmental trajectories as snails reared without predators. Further, the snails possessed wide developmental windows for inducible defenses that extended past sexual maturity. However, being induced later in development appeared to have an associated cost (i.e., decreased shell thickness) that was not found when water bugs were always present. This epiphenotype (i.e., new shell formation as an extension of the current shell) suggests that resource limitation plays an important role in responses to temporal variation in predation risk and may have critical ecological costs that limit the benefits of the inducible defense. Lastly, the ability of snails to completely reverse their defenses was limited to early in ontogeny due to the constraints associated with modular growth of shell material. In sum, we demonstrate that taking a developmental perspective is extremely valuable for understanding the ecology of inducible defenses.     

Effects of belowground biota on primary and secondary metabolites in Brassica oleracea

Summary. Soil organisms in direct and indirect interaction with plant roots affect aboveground herbivores, likely by inducing different plant responses. We investigated the combined effects of the root-knot nematode Meloidogyne incognita (in direct interaction with roots) and the endogeic earthworm Octolasion tyrtaeum (in indirect interaction with roots) on the performance of Brassica oleracea. Both earthworms and nematodes increased N uptake and shoot biomass of B. oleracea. Earthworm activity mobilized more soil N than litter N, and herbivory by nematodes tended to increase the microbial biomass in soil. Only the structural class of sulphur containing glucosinolates was affected by the soil organisms. Earthworms decreased glucoiberin concentrations in B. oleracea shoots. Glucoraphanin was affected by an interaction between earthworms and nematodes.     

Antagonistic interactions between plant competition and insect herbivory

Interspecific competition between plants and herbivory by specialized insects can have synergistic effects on the growth and performance of the attacked host plant. We tested the hypothesis that competition between plants may also negatively affect the performance of herbivores as well as their top-down effect on the host plant. In such a case, the combined effects of competition and herbivory may be less than expected from a simple multiplicative response. In other words, competition and herbivory may interact antagonistically. In a greenhouse experiment, Poa annua was grown in the presence or absence of a competitor (either Plantago lanceolata or Trifolium repens), as well as with or without a Poa-specialist aphid herbivore. Both competition and herbivory negatively affected Poa growth. Competition also reduced aphid density on Poa. This effect could in part be explained by changes in the biomass and the nitrogen content of Poa shoots. In treatments with competitors, reduced aphid densities alleviated the negative effect of herbivory on above- and belowground Poa biomass. Hence, we were able to demonstrate an antagonistic interaction between plant-plant interspecific competition and herbivory. However, response indices suggested that antagonistic interactions between competition and herbivory were contingent on the identity of the competitor. We found the antagonistic effect only in treatments with T. repens as the competitor. We conclude that both competitor identity and the herbivore's ability to respond with changes in its density or activity to plant competition affect the magnitude and direction (synergistic vs. antagonistic) of the interaction between competition and herbivory on plant growth.     

Untangling positive and negative biotic interactions: views from above and below ground in a forest ecosystem

In ecological communities, the outcome of plant-plant interactions represents the net effect of positive and negative interactions occurring above and below ground. Untangling these complex relationships can provide a better understanding of mechanisms that underlie plant-plant interactions and enhance our ability to predict population, community, and ecosystem effects of biotic interactions. In forested ecosystems, tree seedlings interact with established vegetation, but the mechanisms and outcomes of these interactions are not well understood. To explore such mechanisms, we manipulated above- and belowground interactions among tree seedlings, shrubs, and trees and monitored seedling survival and growth of six species (Pinus banksiana, Betula papyrifera, P. resinosa, Quercus rubra, P. strobus, and Acer rubrum) in mature pine-dominated forest in northern Minnesota, USA. The forest had a moderately open canopy and sandy soils. Understory manipulations were implemented in the forest interior and in large gaps and included removal of shrubs (no interactions), tieback of shrubs (belowground), removal of shrubs with addition of shade (aboveground), and unmanipulated shrubs (both below- and aboveground). We found that shrubs either suppressed or facilitated seedling survival and growth depending on the seedling species, source of interaction (e.g., above- or belowground), and ecological context (e.g., gap or forest interior). In general, shrubs strongly influenced survival and growth in gaps, with more modest effects in the forest interior. In gaps, the presence of shrub roots markedly decreased seedling growth and survival, supporting the idea that belowground competition may be more important in dry, nutrient-poor sites. Shrub shade effects were neutral for three species and facilitative for the other three. Facilitation was more likely for shade-tolerant species. In the forest interior, shrub shade negatively affected seedling survival for the most shade-intolerant species. For several species the net effect of shrubs masked the existence of both positive and negative interactions above and below ground. Our results highlight the complexity of plant-plant interactions, demonstrate that outcomes of these interactions vary with the nature of resource limitation and the ecophysiology of the species involved, and suggest that ecological theory that rests on particular notions of plant-plant interactions (e.g., competition) should consider simultaneous positive and negative interactions occurring above and below ground.