Aboveground Herbivores Research Articles

Responses of belowground communities to large aboveground herbivores: Meta-analysis reveals biome-dependent patterns and critical research gaps.

The importance of herbivore-plant and soil biota-plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta-analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root-feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.

Sequential above- and belowground herbivory modifies plant responses depending on herbivore identity

BackgroundHerbivore-induced changes in plant traits can cause indirect interactions between spatially and/or temporally separated herbivores that share the same host plant. Feeding modes of the herbivores is one of the major factors that influence the outcome of such interactions. Here, we tested whether the effects of transient aboveground herbivory for seven days by herbivores of different feeding guilds on tomato plants (Solanum lycopersicum) alters their interaction with spatially as well as temporally separated belowground herbivores.Results The transient aboveground herbivory by both chewing caterpillars (Spodoptera exigua) and sucking aphids (Myzus persicae) had significant impacts on plant traits such as plant growth, resource allocation and phytohormone contents. While the changes in plant traits did not affect the overall performance of the root-knot nematodes (Meloidogyne incognita) in terms of total number of galls, we found that the consequences of aboveground herbivory for the plants can be altered by the subsequent nematode herbivory. For example, plants that had hosted aphids showed compensatory growth when they were later challenged by nematodes, which was not apparent in plants that had hosted only aphids. In contrast, plants that had been fed by S. exigua larvae did not show such compensatory growth even when challenged by nematodes.ConclusionThe results suggest that the earlier aboveground herbivory can modify plant responses to subsequent herbivores, and such modifications may depend upon identity and/or feeding modes of the aboveground herbivores.

Above- and belowground insect herbivory modifies the response of a grassland plant community to nitrogen eutrophication.

Understanding the role that species interactions play in determining the rate and direction of ecosystem change due to nitrogen (N) eutrophication is important for predicting the consequences of global change. Insects might play a major role in this context. They consume substantial amounts of plant biomass and can alter competitive interactions among plants, indirectly shaping plant community composition. Nitrogen eutrophication affects plant communities globally, but there is limited experimental evidence of how insect herbivory modifies plant community response to raised N levels. Even less is known about the roles of above- and belowground herbivory in shaping plant communities, and how the interaction between the two might modify a plant community's response to N eutrophication. We conducted a 3-yr field experiment where grassland plant communities were subjected to above- and belowground insect herbivory with and without N addition, in a full-factorial design. We found that herbivory modified plant community responses to N addition. Aboveground herbivory decreased aboveground plant community biomass by 21%, but only at elevated N. When combined, above- and belowground herbivory had a stronger negative effect on plant community biomass at ambient N (11% decrease) than at elevated N (4% decrease). In addition, herbivory shifted the functional composition of the plant community, and the magnitude of the shifts depended on the N level. The N and herbivory treatments synergistically conferred a competitive advantage to forbs, which benefited when both herbivory types were present at elevated N. Evenness among the plant species groups increased when aboveground herbivory was present, but N addition attenuated this increase. Our results demonstrate that a deeper understanding of how plant-herbivore interactions above and below ground shape the composition of a plant community is crucial for making reliable predictions about the ecological consequences of global change.

Responses of soil biota and nitrogen availability to an invasive plant under aboveground herbivory

Aims Recently, much attention has been paid to the plant-mediated effects of aboveground herbivory on soil ecosystems. However, studies about the herbivore-induced effects of invasive plants on soil ecosystem are still lacking. In this study, we aimed to examine the soil biota and nutrient availability to an invasive plant under aboveground herbivory stress, and compare the soil responses with a native plant.

Herbivores sculpt leaf traits differently in grasslands depending on life form and land-use histories.

Vertebrate and invertebrate herbivores alter plant communities directly by selectively consuming plant species; and indirectly by inducing morphological and physiological changes to plant traits that provide competitive or survivorship advantages to some life forms over others. Progressively excluding aboveground herbivore communities (ungulates, medium and small sized mammals, invertebrates) over five growing seasons, we explored how leaf morphology (specific leaf area or SLA) and nutrition (nitrogen, carbon, phosphorous, potassium, sodium, and calcium) of different plant life forms (forbs, legumes, grasses, sedges) correlated with their dominance. We experimented in two subalpine grassland types with different land-use histories: (1) heavily grazed, nutrient-rich, short-grass vegetation and (2) lightly grazed, lower nutrient tall-grass vegetation. We found differences in leaf traits between treatments where either all herbivores were excluded or all herbivores were present, showing the importance of considering the impacts of both vertebrates and invertebrates on the leaf traits of plant species. Life forms responses to the progressive exclusion of herbivores were captured by six possible combinations: (1) increased leaf size and resource use efficiency (leaf area/nutrients) where lower nutrient levels are invested in leaf construction, but a reduction in the number of leaves, for example, forbs in both vegetation types, (2) increased leaf size and resource use efficiency, for example, legumes in short grass, (3) increased leaf size but a reduction in the number of leaves, for example, legumes in the tall grass, (4) increased number of leaves produced and increased resource use efficiency, for example, grasses in the short grass, (5) increased resource use efficiency of leaves only, for example, grasses and sedges in the tall grass, and (6) no response in terms of leaf construction or dominance, for example, sedges in the short grass. Although we found multiple possible responses by life forms to progressive exclusion of herbivores, we also found some important generalities. Changes in leaf traits of legumes and grasses correlated with their increasing dominance in the short-grass vegetation and plants were more efficient at constructing photosynthetic tissue when herbivores are present with few exceptions. These results demonstrate that vertebrate and invertebrate herbivores are essential to maintain plant species richness and resource-use efficiency.

An experimental test of the EICA Hypothesis in multiple ranges: invasive populations outperform those from the native range independent of insect herbivore suppression.

The success of invasive plants may reflect environmental differences in their native and introduced ranges including both abiotic and biotic conditions, such as release from aboveground herbivory. However, in response to these novel conditions, plants from invasive populations may have higher growth rates and lower defense levels compared to those in the native range. This may contribute to their success in the introduced range but perhaps not in the native range. Here, we grew 1000 Triadica sebifera plants from 14 native and introduced populations in seven common gardens with unmanaged background vegetation for three growing seasons in three geographic venues that varied in T. sebifera status and insect herbivore communities: Texas -T. sebifera is invasive, low levels of generalist herbivory; Hawaii - T. sebifera introduced but not invasive, high levels of generalist herbivory from exotic herbivores; China - native range, both generalist and specialist herbivores. We suppressed aboveground insects with insecticide on half the plants. Aboveground damage in the first growing season was lowest in Texas and insecticide sprays reduced damage in China. At the end of the first growing season, plants were tallest on average in China and shortest in Hawaii. However, height in later years and mass were highest on average in Texas and lowest in Hawaii. However, there was large variation in damage and plant performance among gardens within venues. Our results suggest that more rapid aboveground growth rates contribute to T. sebifera's success in both the invasive and native ranges independent of aboveground herbivory. However, strong variation among sites indicates that T. sebifera plants from invasive populations only have a strong advantage in a subset of sites in Texas.

Timing of simulated aboveground herbivory influences population dynamics of root-feeding nematodes

AimsPlant damage inflicted by aboveground herbivores can occur at different stages of plant development and can induce plant responses that affect the growth of belowground herbivores. This study explores impacts of aboveground herbivory at different plant development stages on the population dynamics of root-feeding nematodes.MethodsWe simulated aboveground herbivory by clipping the foliage of the grass species Holcus lanatus, and tested how plant defoliation at different times (1, 4 or 7 weeks after nematode inoculation) influenced the population of two root-feeding nematode species: the endoparasitic Pratylenchus penetrans and the ectoparasitic Tylenchorhynchus dubius.ResultsDefoliation increased the total abundance of P. penetrans and the number per unit root mass (density) of both P. penetrans and T. dubius. Defoliation enhanced the density of P. penetrans, however, only when plants were defoliated early. Timing did not influence the density of T. dubius, although both abundance and density increased over time. Defoliation increased the nitrogen concentration of plant roots, but reduced root biomass. The strongest reduction of root biomass occurred after early defoliation.ConclusionsOur study indicates that plant responses to aboveground herbivory and their effects on belowground herbivores can be influenced by the time when plants are defoliated, as well as by the belowground herbivore species and their interactions.

Mechanisms and ecological implications of plant-mediated interactions between belowground and aboveground insect herbivores

Plant-mediated interactions between belowground (BG) and aboveground (AG) herbivores have received increasing interest recently. However, the molecular mechanisms underlying ecological consequences of BG–AG interactions are not fully clear yet. Herbivore-induced plant defenses are complex and comprise phytohormonal signaling, gene expression and production of defensive compounds (defined here as response levels), each with their own temporal dynamics. Jointly they shape the response that will be expressed. However, because different induction methods are used in different plant-herbivore systems, and only one or two response levels are measured in each study, our ability to construct a general framework for BG–AG interactions remains limited. Here we aim to link the mechanisms to the ecological consequences of plant-mediated interactions between BG and AG insect herbivores. We first outline the molecular mechanisms of herbivore-induced responses involved in BG–AG interactions. Then we synthesize the literature on BG–AG interactions in two well-studied plant-herbivore systems, Brassica spp. and Zea mays, to identify general patterns and specific differences. Based on this comprehensive review, we conclude that phytohormones can only partially mimic induction by real herbivores. BG herbivory induces resistance to AG herbivores in both systems, but only in maize this involves drought stress responses. This may be due to morphological and physiological differences between monocotyledonous (maize) and dicotyledonous (Brassica) species, and differences in the feeding strategies of the herbivores used. Therefore, we strongly recommend that future studies explicitly account for these basic differences in plant morphology and include additional herbivores while investigating all response levels involved in BG–AG interactions.

Eliciting maize defense pathways aboveground attracts belowground biocontrol agents.

Plant defense pathways mediate multitrophic interactions above and belowground. Understanding the effects of these pathways on pests and natural enemies above and belowground holds great potential for designing effective control strategies. Here we investigate the effects of aboveground stimulation of plant defense pathways on the interactions between corn, the aboveground herbivore adult Diabrotica speciosa, the belowground herbivore larval D. speciosa, and the subterranean ento-mopathogenic nematode natural enemy Heterorhabditis amazonensis. We show that adult D. speciosa recruit to aboveground herbivory and methyl salicylate treatment, that larval D. speciosa are relatively indiscriminate, and that H. amazonensis en-tomopathogenic nematodes recruit to corn fed upon by adult D. speciosa. These results suggest that entomopathogenicnematodes belowground can be highly attuned to changes in the aboveground parts of plants and that biological control can be enhanced with induced plant defense in this and similar systems.

Tomato belowground–aboveground interactions: Rhizophagus irregularis affects foraging behavior and life history traits of the predator Macrolophus pygmaeus (Hemiptera: Miridae)

In recent years, studies on arbuscular mycorrhizal fungi (AMF) have been revealing that the belowground symbiosis can influence the performance of aboveground herbivores and their natural enemies through its effects on the host plant. In this study, we tested whether the colonization of tomato plants by the arbuscular mycorrhizal fungus Rhizophagus irregularis (Syn. Glomus intraradices Schenk and Smith) (Glomeromycota: Glomeraceae) affects the performance of the zoophytophagous mirid bug Macrolophus pygmaeus Rambur (Hemiptera: Miridae). Mycorrhizal colonization in tomato plants positively influenced the predator host-plant acceptance for feeding and oviposition, as well as nymphal survival and female weight. We hypothesize that AMF can modify mirid bug foraging behavior and performance.

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.

Plant genotypes affect aboveground and belowground herbivore interactions by changing chemical defense.

Spatially separated aboveground (AG) and belowground (BG) herbivores are closely linked through shared host plants, and both patterns of AG-BG interactions and plant responses may vary among plant genotypes. We subjected invasive (USA) and native (China) genotypes of tallow tree (Triadica sebifera) to herbivory by the AG specialist leaf-rolling weevil Heterapoderopsis bicallosicollis and/or the root-feeding larvae of flea beetle Bikasha collaris. We measured leaf damage and leaves rolled by weevils, quantified beetle survival, and analyzed flavonoid and tannin concentrations in leaves and roots. AG and BG herbivores formed negative feedbacks on both native and invasive genotypes. Leaf damage by weevils and the number of beetle larvae emerging as adults were higher on invasive genotypes. Beetles reduced weevil damage and weevils reduced beetle larval emergence more strongly for invasive genotypes. Invasive genotypes had lower leaf and root tannins than native genotypes. BG beetles decreased leaf tannins of native genotypes but increased root tannins of invasive genotypes. AG herbivory increased root flavonoids of invasive genotypes while BG herbivory decreased leaf flavonoids. Invasive genotypes had lower AG and BG herbivore resistance, and negative AG-BG herbivore feedbacks were much stronger for invasive genotypes. Lower tannin concentrations explained overall better AG and BG herbivore performances on invasive genotypes. However, changes in tannins and flavonoids affected AG and BG herbivores differently. These results suggest that divergent selection on chemical production in invasive plants may be critical in regulating herbivore performances and novel AG and BG herbivore communities in new environments.

Aboveground mammal and invertebrate exclusions cause consistent changes in soil food webs of two subalpine grassland types, but mechanisms are system‐specific

Ungulates, smaller mammals, and invertebrates can each affect soil biota through their influence on vegetation and soil characteristics. However, direct and indirect effects of the aboveground biota on soil food webs remain to be unraveled. We assessed effects of progressively excluding aboveground large‐, medium‐ and small‐sized mammals as well as invertebrates on soil nematode diversity and feeding type abundances in two subalpine grassland types: short‐ and tall‐grass vegetation. We explored pathways that link exclusions of aboveground biota to nematode feeding type abundances via changes in plants, soil environment, soil microbial biomass, and soil nutrients.In both vegetation types, exclusions caused a similar shift toward higher abundance of all nematode feeding types, except plant feeders, lower Shannon diversity, and lower evenness. These effects were strongest when small mammals, or both small mammals and invertebrates were excluded in addition to excluding larger mammals. Exclusions resulted in a changed abiotic soil environment that only affected nematodes in the short‐grass vegetation. In each vegetation type, exclusion effects on nematode abundances were mediated by different drivers related to plant quantity and quality. In the short‐grass vegetation, not all exclusion effects on omni–carnivorous nematodes were mediated by the abundance of lower trophic level nematodes, suggesting that omni–carnivores also depended on other prey than nematodes.We conclude that small aboveground herbivores have major impacts on the soil food web of subalpine short‐ and tall‐grass ecosystems. Excluding aboveground animals caused similar shifts in soil nematode assemblages in both subalpine vegetation types, however, mechanisms turned out to be system‐specific.

Effects of aboveground herbivory on plants with long-term belowground biomass storage

Plant tolerance to herbivory is contingent on multiple traits and adaptive mechanisms, which makes it a complex response with ecological implications. In plants with long-term belowground storage, allocation of biomass to inaccessible parts belowground in response to folivory is a well-recognized tolerance mechanism. In temperate regions, spring growth from buried rootstock is common among winter deciduous plants and is often followed by regrowth after defoliation, both of which draws resources from the stored reserves. We developed a mathematical model to analyze this tolerance response in a winter deciduous plant with long-term belowground biomass when it is defoliated by a specialist insect folivore. The model explores how three closely associated traits—(1) belowground biomass allocation to roots, (2) spring utilization of stored reserves, and (3) post-defoliation regrowth capacity—modulate the persistence and dynamics of the plant and herbivore populations. Model results show that allocation to belowground storage is not only a critical component of tolerance but also influences the herbivore population dynamics in ways that depend on how and when plant biomass is allocated and used. Low belowground biomass allocation and high storage utilization combined with poor photosynthetic growth caused extirpation of the plant population by the defoliating insects. Stable coexistence of the plant at low biomass along with its specialist insect required a moderate amount of post-herbivory belowground allocation. High values of belowground biomass allocation, storage utilization, and photosynthetic growth resulted in sustained cycles of the herbivore and plant populations. Interestingly, utilization of stored reserves had conflicting influence on above and belowground biomass, and strongly affected herbivore population dynamics. Our model thus highlights the complexity of tolerance response when it involves multiple traits and mechanisms as evinced by winter deciduous plants. We close by discussing the implications of our findings for the contributions of defoliating insects to biocontrol programs.

Potato tuber herbivory increases resistance to aboveground lepidopteran herbivores.

Plants mediate interactions between aboveground and belowground herbivores. Although effects of root herbivory on foliar herbivores have been documented in several plant species, interactions between tuber-feeding herbivores and foliar herbivores are rarely investigated. We report that localized tuber damage by Tecia solanivora (Guatemalan tuber moth) larvae reduced aboveground Spodoptera exigua (beet armyworm) and Spodoptera frugiperda (fall armyworm) performance on Solanum tuberosum (potato). Conversely, S. exigua leaf damage had no noticeable effect on belowground T. solanivora performance. Tuber infestation by T. solanivora induced systemic plant defenses and elevated resistance to aboveground herbivores. Lipoxygenase 3 (Lox3), which contributes to the synthesis of plant defense signaling molecules, had higher transcript abundance in T. solanivora-infested leaves and tubers than in equivalent control samples. Foliar expression of the hydroxycinnamoyl-CoA quinate hydroxycinnamoyl transferase (HQT) and 3-hydroxy-3-methylglutaryl CoA reductase I (HMGR1) genes, which are involved in chlorogenic acid and steroidal glycoalkaloid biosynthesis,respectively, also increased in response to tuber herbivory. Leaf metabolite profiling demonstrated the accumulation of unknown metabolites as well as the known potato defense compounds chlorogenic acid, α-solanine, and α-chaconine. When added to insect diet at concentrations similar to those found in potato leaves, chlorogenic acid, α-solanine, and α-chaconine all reduced S. exigua larval growth. Thus, despite the fact that tubers are a metabolic sink tissue, T. solanivora feeding elicits a systemic signal that induces aboveground resistance against S. exigua and S. frugiperda by increasing foliar abundance of defensive metabolites.

Variation in herbivory along a latitudinal gradient for native and exotic Asteraceae

It has long been hypothesized that biotic interactions, including herbivory, are most intense at lower latitudes. However, this generalization has recently been challenged with studies showing that latitudinal gradients in damage may be rarer than previously believed. Additionally, most studies have focused on herbivory of native species, so it remains unknown whether natives and exotics follow similar patterns. This study compares rates of aboveground herbivory of multiple native and non-native Asteraceae across a latitudinal gradient, with a more detailed investigation of a focal exotic, Cirsium arvense. Herbivory of multiple tissue types was quantified for all species across an 815 km transect in Ontario, Canada. The native Asteraceae included in the survey typically experienced a decline in folivory with increasing latitude. Herbivory patterns for the exotic species were less clear; while most experienced high damage at the southernmost site, some also experienced high damage rates at mid-latitudes. For the focal species C. arvense, leaf and stem herbivory declined with increasing latitude, although seed damage showed strong regional variation across the invaded range. These results show that latitudinal variation in herbivory is highly dependent on the plant species being investigated, the tissue type being measured, and the type of herbivore(s) causing the damage. In some cases, populations in marginal areas might benefit from reduced damage by some groups of herbivores. In other cases, factors such as the availability of suitable habitat, the biology of specific enemies, and the origin of the host plant may override the influence of latitude on host performance.

Aboveground insect herbivory increases plant competitive asymmetry, while belowground herbivory mitigates the effect.

Insect herbivores can shift the composition of a plant community, but the mechanism underlying such shifts remains largely unexplored. A possibility is that insects alter the competitive symmetry between plant species. The effect of herbivory on competition likely depends on whether the plants are subjected to aboveground or belowground herbivory or both, and also depends on soil nitrogen levels. It is unclear how these biotic and abiotic factors interactively affect competition. In a greenhouse experiment, we measured competition between two coexisting grass species that respond differently to nitrogen deposition: Dactylis glomerata L., which is competitively favoured by nitrogen addition, and Festuca rubra L., which is competitively favoured on nitrogen-poor soils. We predicted: (1) that aboveground herbivory would reduce competitive asymmetry at high soil nitrogen by reducing the competitive advantage of D. glomerata; and (2), that belowground herbivory would relax competition at low soil nitrogen, by reducing the competitive advantage of F. rubra. Aboveground herbivory caused a 46% decrease in the competitive ability of F. rubra, and a 23% increase in that of D. glomerata, thus increasing competitive asymmetry, independently of soil nitrogen level. Belowground herbivory did not affect competitive symmetry, but the combined influence of above- and belowground herbivory was weaker than predicted from their individual effects. Belowground herbivory thus mitigated the increased competitive asymmetry caused by aboveground herbivory. D. glomerata remained competitively dominant after the cessation of aboveground herbivory, showing that the influence of herbivory continued beyond the feeding period. We showed that insect herbivory can strongly influence plant competitive interactions. In our experimental plant community, aboveground insect herbivory increased the risk of competitive exclusion of F. rubra. Belowground herbivory appeared to mitigate the influence of aboveground herbivory, and this mechanism may play a role for plant species coexistence.

Soil food web stability in response to grazing in a semi-arid prairie: The importance of soil textural heterogeneity

Grazing of grasslands by large herbivores is a form of land use intensification that affects not only plant communities but also soil biota and the ecosystem services that it provides. While grassland ecosystem responses to grazing have been extensively studied, few studies have focused on the effects of aboveground herbivores on belowground diversity and functions. In this work, we quantified effects of grazing on the structure, function and dynamic stability of soil food webs. We sampled a long-term grazing manipulation in a semi-arid shortgrass steppe (USA Great Plains) at sites showing contrasting soil textures. Treatments included native steppe plots that have been moderately grazed since 1939 paired with plots totally protected from grazing since 1996. We sampled our plots for soil C and N, and for soil biota, separated microbes and micro- and mesofauna in trophic functional groups and defined trophic relationships. We used models to estimate carbon and nitrogen mineralization, energy flow throughout the food web, interaction strengths between trophic groups at steady-state and, eventually, asymptotic (near-equilibrium or local) stability (Moore and de Ruiter, 2012). Soil food web response to grazing depended on soil texture and organic matter content. In our food webs, most energy flowed through the fungal and bacterial detritus-based channels (sensu Moore and Hunt, 1988). There was a clear asymmetry between the amount of energy flowing through each of the two channels and, the higher this asymmetry, the higher was food web stability. Stability was affected by both grazing and soil properties (increased under grazing in high clay soils with high organic matter content but decreased in less organic loam sandy soils), and positively associated with soil organic matter content. Overall, we found that the carbon flow through the soil food web of the shortgrass steppe is responsive to grazing in ways that altered stability and that structural, functional, and dynamic attributes are sensitive parameters for evaluating soil response to land use under changing scenarios.

Root-Lesion Nematodes Suppress Cabbage Aphid Population Development by Reducing Aphid Daily Reproduction

Empirical studies have shown that belowground feeding herbivores can affect the performance of aboveground herbivores in different ways. Often the critical life-history parameters underlying the observed performance effects remain unexplored. In order to better understand the cause for the observed effects on aboveground herbivores, these ecological mechanisms must be better understood. In this study we combined empirical experiments with a modeling approach to analyze the effect of two root feeding endoparasitic nematodes with different feeding strategies on the population growth of the aboveground feeding specialist aphid Brevicoryne brassicae on Brassica nigra. The aim was to test whether emerging differences in life history characteristics (days until reproduction, daily reproduction) would be sufficient to explain observed differences in aphid population development on plants with and without two species of nematodes. Aphid numbers were lower on plants with Pratylenchus penetrans in comparison to aphid numbers on plants with Meloidogyne spp. A dedicated experiment showed that aphid daily reproduction was lower on plants with P. penetrans (3.08 offspring female–1 day–1) in comparison to both uninfested plants and plants with Meloidogyne spp. (3.50 offspring female–1 day–1). The species-specific reduction of aphid reproduction appeared independent of changes in amino acids, soluble sugars or the glucosinolate sinigrin in the phloem. An individual-based model revealed that relatively small differences in reproduction rate per female were sufficient to yield a similar difference in aphid populations as was found in the empirical experiments.

Roots under attack: contrasting plant responses to below- and aboveground insect herbivory.

The distinctive ecology of root herbivores, the complexity and diversity of root-microbe interactions, and the physical nature of the soil matrix mean that plant responses to root herbivory extrapolate poorly from our understanding of responses to aboveground herbivores. For example, root attack induces different changes in phytohormones to those in damaged leaves, including a lower but more potent burst of jasmonates in several plant species. Root secondary metabolite responses also differ markedly, although patterns between roots and shoots are harder to discern. Root defences must therefore be investigated in their own ecophysiological and evolutionary context, specifically one which incorporates root microbial symbionts and antagonists, if we are to better understand the battle between plants and their hidden herbivores.