Root volatiles in plant–plant interactions II: Root volatiles alter root chemistry and plant–herbivore interactions of neighbouring plants

Volatile organic compounds (VOCs) emitted by plant roots can influence the germination and growth of neighbouring plants. However, little is known about the effects of root VOCs on plant–herbivore interactions of neighbouring plants. The spotted knapweed (Centaurea stoebe) constitutively releases high amounts of sesquiterpenes into the rhizosphere. Here, we examine the impact of C. stoebe root VOCs on the primary and secondary metabolites of sympatric Taraxacum officinale plants and the resulting plant‐mediated effects on a generalist root herbivore, the white grub Melolontha melolontha. We show that exposure of T. officinale to C.stoebe root VOCs does not affect the accumulation of defensive secondary metabolites but modulates carbohydrate and total protein levels in T. officinale roots. Furthermore, VOC exposure increases M. melolontha growth on T. officinale plants. Exposure of T. officinale to a major C. stoebe root VOC, the sesquiterpene (E)‐β‐caryophyllene, partially mimics the effect of the full root VOC blend on M. melolontha growth. Thus, releasing root VOCs can modify plant–herbivore interactions of neighbouring plants. The release of VOCs to increase the susceptibility of other plants may be a form of plant offense.     

Direct and indirect effects of episodic frost on plant growth and reproduction in subalpine wildflowers

Frost is an important episodic event that damages plant tissues through the formation of ice crystals at or below freezing temperatures. In montane regions, where climate change is expected to cause earlier snow melt but may not change the last frost‐free day of the year, plants that bud earlier might be directly impacted by frost through damage to flower buds and reproductive structures. However, the indirect effects of frost mediated through changes in plant–pollinator interactions have rarely been explored. We examined the direct and pollinator‐mediated indirect effects of frost on three wildflower species in southwestern Colorado, USA, Delphinium barbeyi (Ranunculaceae), Erigeron speciosus (Asteraceae), and Polemonium foliosissimum (Polemoniaceae), by simulating moderate (?1 to ?5°C) frost events in early spring in plants in situ. Subsequently, we measured plant growth, and upon flowering measured flower morphology and phenology. Throughout the flowering season, we monitored pollinator visitation and collected seeds to measure plant reproduction. We found that frost had species‐specific direct and indirect effects. Frost had direct effects on two of the three species. Frost significantly reduced flower size, total flowers produced, and seed production of Erigeron. Furthermore, frost reduced aboveground plant survival and seed production for Polemonium. However, we found no direct effects of frost on Delphinium. When we considered the indirect impacts of frost mediated through changes in pollinator visitation, one species, Erigeron, incurred indirect, negative effects of frost on plant reproduction through changes in floral traits and pollinator visitation, along with direct effects. Overall, we found that flowering plants exhibited species‐specific direct and pollinator‐mediated indirect responses to frost, thus suggesting that frost may play an important role in affecting plant communities under climate change.     

Soil conditioning effects of Phragmites australis on native wetland plant seedling survival

Interactions between introduced plants and soils they colonize are central to invasive species success in many systems. Belowground biotic and abiotic changes can influence the success of introduced species as well as their native competitors. All plants alter soil properties after colonization but, in the case of many invasive plant species, it is unclear whether the strength and direction of these soil conditioning effects are due to plant traits, plant origin, or local population characteristics and site conditions in the invaded range. Phragmites australis in North America exists as a mix of populations of different evolutionary origin. Populations of endemic native Phragmites australis americanus are declining, while introduced European populations are important wetland invaders. We assessed soil conditioning effects of native and non‐native P. australis populations on early and late seedling survival of native and introduced wetland plants. We further used a soil biocide treatment to assess the role of soil fungi on seedling survival. Survival of seedlings in soils colonized by P. australis was either unaffected or negatively affected; no species showed improved survival in P. australis‐conditioned soils. Population of P. australis was a significant factor explaining the response of seedlings, but origin (native or non‐native) was not a significant factor. Synthesis: Our results highlight the importance of phylogenetic control when assessing impacts of invasive species to avoid conflating general plant traits with mechanisms of invasive success. Both native (noninvasive) and non‐native (invasive) P. australis populations reduced seedling survival of competing plant species. Because soil legacy effects of native and non‐native P. australis are similar, this study suggests that the close phylogenetic relationship between the two populations, and not the invasive status of introduced P. australis, is more relevant to their soil‐mediated impact on other plant species.     

Ecosystem nitrogen fixation throughout the snow‐free period in subarctic tundra: effects of willow and birch litter addition and warming

Nitrogen (N) fixation in moss‐associated cyanobacteria is one of the main sources of available N for N‐limited ecosystems such as subarctic tundra. Yet, N₂ fixation in mosses is strongly influenced by soil moisture and temperature. Thus, temporal scaling up of low‐frequency in situ measurements to several weeks, months or even the entire growing season without taking into account changes in abiotic conditions cannot capture the variation in moss‐associated N₂ fixation. We therefore aimed to estimate moss‐associated N₂ fixation throughout the snow‐free period in subarctic tundra in field experiments simulating climate change: willow (Salix myrsinifolia) and birch (Betula pubescens spp. tortuosa) litter addition, and warming. To achieve this, we established relationships between measured in situ N₂ fixation rates and soil moisture and soil temperature and used high‐resolution measurements of soil moisture and soil temperature (hourly from May to October) to model N₂ fixation. The modelled N₂ fixation rates were highest in the warmed (2.8 ± 0.3 kg N ha^?1) and birch litter addition plots (2.8 ± 0.2 kg N ha^?1), and lowest in the plots receiving willow litter (1.6 ± 0.2 kg N ha^?1). The control plots had intermediate rates (2.2 ± 0.2 kg N ha^?1). Further, N₂ fixation was highest during the summer in the warmed plots, but was lowest in the litter addition plots during the same period. The temperature and moisture dependence of N₂ fixation was different between the climate change treatments, indicating a shift in the N₂ fixer community. Our findings, using a combined empirical and modelling approach, suggest that a longer snow‐free period and increased temperatures in a future climate will likely lead to higher N₂ fixation rates in mosses. Yet, the consequences of increased litter fall on moss‐associated N₂ fixation due to shrub expansion in the Arctic will depend on the shrub species’ litter traits.     

Entrapped sand as a plant defence: effects on herbivore performance and preference

1. Abrasive material in the diet of herbivorous organisms comes from a variety of sources, including crystalline silica or calcium in plant tissues, accidentally ingested soil while digging or grazing, and entrapped substrate on the surfaces of plants. A wide variety of plants entrap substrate, usually with glandular trichomes. 2. A previous study demonstrated that entrapped sand provided resistance to herbivory in the field. In this study, the following questions were addressed: how does entrapped sand on Abronia latifolia (Nyctaginaceae) leaves and stems affect preference and performance of a common herbivore, the large‐bodied caterpillar Hyles lineata (Sphingidae); does this effect differ from those experienced by an internally feeding leaf miner? 3. Using a combination of experimental and observational approaches, it was found that sand comprised ～4–5% of ingested weight during normal feeding of H. lineata caterpillars. This entrapped sand caused extensive wear to their mandibles, they avoided sand‐covered plants when given the choice, and the sand negatively impacted performance metrics, including pupal weight, development time, and growth rate. In contrast, a leaf‐mining caterpillar did not have a preference for or against feeding on sandy plants. 4. These results are similar to studies on mandibular wear due to grasses, and herbivorous insects that feed on these two plant groups may have similar morphologies. It is hypothesised that increased wear potential may be a convergent solution to abrasive plants in both mammals (hypsodonty) and insects.     

Wildlife species benefitting from a greener Arctic are most sensitive to shrub cover at leading range edges

Widespread expansion of shrubs is occurring across the Arctic. Shrub expansion will substantially alter arctic wildlife habitats. Identifying which wildlife species are most affected by shrubification is central to predicting future arctic community composition. Through meta‐analysis, we synthesized the published evidence for effects of canopy‐forming shrubs on birds and mammals in the Arctic and Subarctic. We examined variation in species behaviour, distribution and population dynamics in birds and mammals in response to shrub cover (including shrub cover indicators such as shrub occurrence, extent, density and height). We also assessed the degree of heterogeneity in wildlife responses to shrub cover and synthesized the remaining literature that did not fit the criteria for our quantitative meta‐analyses. Species from higher green vegetation biomass habitats (high Normalized Difference Vegetation Index, NDVI, across their distribution) were more likely to respond positively to shrub cover, demonstrating the potential for species to expand from boreal to arctic habitats under shrubification. Wildlife populations located in the lowest vegetation biomass (low NDVI) areas of their species’ range had the greatest proportion of positive responses to shrub cover, highlighting how increases in performance at leading edges of invaders distributions may be particularly rapid. This demonstrates the need to study species at these leading edges to accurately predict expansion potential. Arctic specialists were poorly represented across studies (limited to 5 bird and 0 mammal species), this knowledge gap potentially explains the few reported negative effects of shrub cover (3 of 29 species). Species responses to shrub cover showed substantial heterogeneity and varied among sites and years in all studies with sufficient replication to detect such variation. Our study highlights the importance of responses at species range edges in determining outcomes of shrubification for arctic birds and mammals and the need for greater examination of potential wildlife losers under shrubification.     

Genomic evidence of genetic variation with pleiotropic effects on caterpillar fitness and plant traits in a model legume

Plant–insect interactions are ubiquitous, and have been studied intensely because of their relevance to damage and pollination in agricultural plants, and to the ecology and evolution of biodiversity. Variation within species can affect the outcome of these interactions. Specific genes and chemicals that mediate these interactions have been identified, but genome‐ or metabolome‐scale studies might be necessary to better understand the ecological and evolutionary consequences of intraspecific variation for plant–insect interactions. Here, we present such a study. Specifically, we assess the consequences of genome‐wide genetic variation in the model plant Medicago truncatula for Lycaeides melissa caterpillar growth and survival (larval performance). Using a rearing experiment and a whole‐genome SNP data set (>5 million SNPs), we found that polygenic variation in M. truncatula explains 9%–41% of the observed variation in caterpillar growth and survival. Genetic correlations among caterpillar performance and other plant traits, including structural defences and some anonymous chemical features, suggest that multiple M. truncatula alleles have pleiotropic effects on plant traits and caterpillar performance (or that substantial linkage disequilibrium exists among distinct loci affecting subsets of these traits). A moderate proportion of the genetic effect of M. truncatula alleles on L. melissa performance can be explained by the effect of these alleles on the plant traits we measured, especially leaf toughness. Taken together, our results show that intraspecific genetic variation in M. truncatula has a substantial effect on the successful development of L. melissa caterpillars (i.e., on a plant–insect interaction), and further point toward traits potentially mediating this genetic effect.     

Nurse species and indirect facilitation through grazing drive plant community functional traits in tropical alpine peatlands

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant–plant–herbivore interactions in stressful environments. We set up a 2‐year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion‐forming plants (hereafter referred to as “nurse” species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the “grazer–nurse–beneficiary” interaction module can be more complex than expected when evaluated in the field.     

Relatedness with plant species in native community influences ecological consequences of range expansions

Global warming is enabling many plant species to expand their range to higher latitudes and altitudes, where they may suffer less from natural aboveground and belowground enemies. Reduced control by natural enemies can enable climate warming‐induced range expanders to gain an advantage in competition with natives and become disproportionally abundant in their new range. However, so far studies have only examined individual growth of range expanders, which have common congeneric plant species in their new range. Thus it is not known how general is this reduced effect of above‐ and belowground enemies and how it operates in communities, where multiple plant species also interact with each other. Here we show that range‐expanding plant species with and without congenerics in the invaded habitats differ in their ecological interactions in the new range. In a community‐level experiment, range‐expanding plant species, both with and without congenerics, suppressed the growth of a herbivore. However, only range expanders without congenerics reduced biomass production of the native plant species. In the present study, range expanders without congenerics allocated more biomass aboveground compared to native plant species, which can explain their competitive advantage. Competitive interaction and also biomass allocation of native plants and their congeneric range expanders were similar. Our results highlight that information about species phylogenetic relatedness with native flora can be crucial for improving predictions about the consequences of climate warming‐induced range expansions.     

Anthropogenic increase in carbon dioxide modifies plant–insect interactions

Industrialisation has elevated atmospheric levels of CO₂ from original 280 ppm to current levels at 400 ppm, which is estimated to double by 2050. Although high atmospheric CO₂ levels affect insect interactions with host plants, the impact of global change on plant defences in response to insect attack is not completely understood. Recent studies have made advances in elucidating the mechanisms of the effects of high CO₂ levels in plant–insect interactions. New studies have proposed that gene regulation and phytohormones regulate resource allocation from photosynthesis to plant defences against insects. Biochemical and molecular studies demonstrated that both defensive hormones jasmonic acid (JA) and ethylene (ET) participate in modulating chemical defences against herbivores in plants grown under elevated CO₂ atmosphere rather than changes in C:N ratio. High atmospheric CO₂ levels increase vulnerability to insect damage by down‐regulating both inducive and constitutive chemical defences regulated by JA and ET. However, elevated CO₂ levels increase the JA antagonistic hormone salicylic acid that increases other chemical defences. How plants grown under elevated CO₂ environment allocate primary metabolites from photosynthesis to secondary metabolism would help to understand innate defences and prevent future herbivory in field crops. We present evidence demonstrating that changes in chemical defences in plants grown under elevated CO₂ environment are hormonal regulated and reject the C:N hypothesis. In addition, we discuss current knowledge of the mechanisms that regulate plants defences against insects in elevated CO₂ atmospheres.     

Effect of plant root symbionts on performance of native woody species in competition with an invasive grass in multispecies microcosms

The majority of terrestrial plants form mutualistic associations with arbuscular mycorrhizal fungi (AMF) and rhizobia (i.e., nitrogen‐fixing bacteria). Understanding these associations has important implications for ecological theory and for restoration practice. Here, we tested whether the presence of AMF and rhizobia influences the performance of native woody plants invaded by a non‐native grass in experimental microcosms. We planted eight plant species (i.e., Acacia acuminata, A. microbotrya, Eucalyptus loxophleba subsp. loxophleba, E. astringens, Calothamnus quadrifidus, Callistemon phoeniceus, Hakea lissocarpha and H. prostrata) in microcosms of field‐conditioned soil with and without addition of AMF and rhizobia in a fully factorial experimental design. After seedling establishment, we seeded half the microcosms with an invasive grass Bromus diandrus. We measured shoot and root biomass of native plants and Bromus, and on roots, the percentage colonization by AMF, number of rhizobia‐forming nodules and number of proteaceous root clusters. We found no effect of plant root symbionts or Bromus addition on performance of myrtaceous, and as predicted, proteaceous species as they rely little or not at all on AMF and rhizobia. Soil treatments with AMF and rhizobia had a strong positive effect (i.e., larger biomass) on native legumes (A. microbotrya and A. acuminata). However, the beneficial effect of root symbionts on legumes became negative (i.e., lower biomass and less nodules) if Bromus was present, especially for one legume, i.e., A. acuminata, suggesting a disruptive effect of the invader on the mutualism. We also found a stimulating effect of Bromus on root nodule production in A. microbotrya and AMF colonization in A. acuminata which could be indicative of legumes’ increased resource acquisition requirement, i.e., for nitrogen and phosphorus, respectively, in response to the Bromus addition. We have demonstrated the importance of measuring belowground effects because the aboveground effects gave limited indication of the effects occurring belowground.     

Experimental warming transforms multiple predator effects in a grassland food web

This experimental study tests new theory for multiple predator effects on communities by using warming to alter predator habitat use and hence direct and indirect interactions in a grassland food web containing two dominant spider predator species, a dominant grasshopper herbivore and grass and herb plants. Experimental warming further offers insight into how climate change might alter direct and indirect effects. Under ambient environmental conditions, spiders used habitat in spatially complementary locations. Consistent with predictions, the multiple predator effect on grasshoppers and on plants was the average of the individual predator effects. Warming strengthened the single predator effects. It also caused the spider species to overlap lower in the vegetation canopy. Consistent with predictions, the system was transformed into an intraguild predation system with the consequent extinction of one spider species. The results portend climate caused loss of predator diversity with important consequences for food web structure and function.     

Phenological phases of the host plant shape plant–treehopper interaction networks

相似文献   

Gut microbes of mammalian herbivores facilitate intake of plant toxins

The foraging ecology of mammalian herbivores is strongly shaped by plant secondary compounds (PSCs) that defend plants against herbivory. Conventional wisdom holds that gut microbes facilitate the ingestion of toxic plants; however, this notion lacks empirical evidence. We investigated the gut microbiota of desert woodrats (Neotoma lepida), some populations of which specialise on highly toxic creosote bush (Larrea tridentata). Here, we demonstrate that gut microbes are crucial in allowing herbivores to consume toxic plants. Creosote toxins altered the population structure of the gut microbiome to facilitate an increase in abundance of genes that metabolise toxic compounds. In addition, woodrats were unable to consume creosote toxins after the microbiota was disrupted with antibiotics. Last, ingestion of toxins by naïve hosts was increased through microbial transplants from experienced donors. These results demonstrate that microbes can enhance the ability of hosts to consume PSCs and therefore expand the dietary niche breadth of mammalian herbivores.     

Low interannual precipitation has a greater negative effect than seedling herbivory on the population dynamics of a short‐lived shrub,Schiedea obovata

Climate projections forecast more extreme interannual climate variability over time, with an increase in the severity and duration of extreme drought and rainfall events. Based on bioclimatic envelope models, it is projected that changing precipitation patterns will drastically alter the spatial distributions and density of plants and be a primary driver of biodiversity loss. However, many other underlying mechanisms can impact plant vital rates (i.e., survival, growth, and reproduction) and population dynamics. In this study, we developed a size‐dependent integral projection model (IPM) to evaluate how interannual precipitation and mollusk herbivory influence the dynamics of a Hawaii endemic short‐lived shrub, Schiedea obovata (Caryophyllaceae). Assessing how wet season precipitation effects population dynamics it critical, as it is the timeframe when most of the foliar growth occurs, plants flower and fruit, and seedlings establish. Temporal variation in wet season precipitation had a greater effect than mollusk herbivory on S. obovata population growth rate , and the impact of interannual precipitation on vital rates shifted across plant ontogeny. Furthermore, wet season precipitation influenced multiple vital rates in contrasting ways and the effect of precipitation on the survival of larger vegetative and reproductively mature individuals contributed the most to variation in the population growth rate. Among all combination of wet season precipitation and herbivory intensities, the only scenario that led to a growing population was when high wet precipitation was associated with low herbivory. Our study highlights the importance of evaluating how abiotic factors and plant–consumer interactions influence an organism across its life cycle to fully understand the underpinning mechanisms that structure its spatial and temporal distribution and abundance. Our results also illustrate that for short‐lived species, like S. obovata, seedling herbivory can have less of an effect on the dynamics of plant populations than decreased interannual precipitation.     

Plant‐mediated effects of host plant density on a specialist herbivore of Solanum carolinense

相似文献   

Trees to treehoppers: genetic variation in host plants contributes to variation in the mating signals of a plant‐feeding insect

Community genetics research has demonstrated ‘bottom‐up’ effects of genetic variation within a plant species in shaping the larger community with which it interacts, such as compositions of arthropod faunas. We demonstrate that such cross‐trophic interactions also influence sexually selected traits. We used a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae) to ask whether male mating signals are influenced by host plant genetic variation. We reared a random sample of the treehoppers on potted replicates of a sample of host plant clone lines. We found that treehopper male signals varied according to the clone line on which they developed, showing that genetic variation in host plants affects male treehoppers' behavioural phenotypes. This is the first demonstration of cross‐trophic indirect genetic effects on a sexually selected trait. We discuss how such effects may play an important role in the maintenance of variation and within‐population phenotypic differentiation, thereby promoting evolutionary divergence.