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Understanding the mechanisms that drive community assembly in species‐rich tropical forest remains a fundamental challenge in ecology. Here, we integrated multivariate functional trait dimensions, phylogeny and metabolomics to test fundamental predictions concerning the role of differentiation with respect to abiotic and biotic niche axes in the maintenance of high local diversity of woody plants in the Euphorbiaceae.We measured 40 functional traits related to resource acquisition, photosynthetic capacity, hydraulic efficiency and secondary‐metabolite profiles generated using untargeted metabolomics in all 26 Euphorbiaceae species in a 20‐ha forest dynamics plot in tropical southwestern China. We examined the correlation structure of 40 traits using a trait networking approach. We coupled these traits with variation in soil nutrients, light environment, soil water content and herbivore pressure within the plot to assess niche differentiation in space. We compared phylogenetic signal among multivariate trait dimensions and secondary metabolites to assess niche differentiation in evolutionary time.Network analysis revealed that a small number of traits with high network centrality reflected variation in ecological strategy among the Euphorbiaceae. Using these high‐centrality traits, we observed significant functional turnover along environmental gradients defined by light, soil moisture, soil nutrients and leaf herbivory, respectively. Most resource utilization traits showed significant phylogenetic signal, whereas almost all defensive traits lacked phylogenetic signal, including species similarity with respect to plant secondary metabolites.Synthesis. Our results suggest that resource‐utilization traits and the habitat associations play a significant role in the niche segregation of co‐occurring woody plants in the Euphorbiaceae. Secondary metabolites, however, may enhance diversity at a finer spatial scale by allowing closely related species with similar functional traits to partition biotic niche space within shared habitats in tropical rainforest. [ABSTRACT FROM AUTHOR]

Plant toxins constitute an effective defense against herbivorous animals. However, many herbivores have evolved adaptations to cope with dietary toxins through detoxification, excretion, sequestration, target site insensitivity and/or via behavioral avoidance. While these adaptations are often directly encoded in herbivore genomes, evidence is accumulating that microbial symbionts can reduce the dose of plant toxins by metabolizing or sequestering them prior to absorption by the herbivore. Here, we describe a few well-studied examples to assess such symbiont-mediated detoxification and showcase different approaches that have been used for their analyses. These include: (i) a host phenotypic route in which the symbiotic association is manipulated to reveal host fitness costs upon toxin exposure in the presence/absence of detoxifying symbionts, including function restoration after symbiont re-infection, (ii) a molecular microbiological approach that focuses on the identification and characterization of microbial genes involved in plant toxin metabolism, and (iii) an analytical chemical route that aims to characterize the conversion of the toxin to less harmful metabolites in vivo and link conversion to the activities of a detoxifying symbiont. The advantages and challenges of each approach are discussed, and it is argued that a multi-pronged strategy combining phenotypic, molecular, and chemical evidence is needed to unambiguously demonstrate microbial contributions to plant toxin reduction and the importance of these processes for host fitness. Given the interdisciplinary nature of the topic, we aim to provide a guideline to researchers interested in symbiont-mediated detoxification and hope to encourage future studies that contribute to a more comprehensive and mechanistic understanding of detoxification in herbivores and their symbionts. [ABSTRACT FROM AUTHOR]

Defense against natural enemies constitutes an important driver of herbivore host range evolution in the wild. Populations of the Baltimore checkerspot butterfly, Euphydryas phaeton (Nymphalidae), have recently incorporated an exotic plant, Plantago lanceolata (Plantaginaceae), into their dietary range. To understand the tritrophic consequences of utilizing this exotic host plant, we examined immune performance, chemical defense, and interactions with a natural entomopathogen (Junonia coenia densovirus, Parvoviridae) across wild populations of this specialist herbivore. We measured three immune parameters, sequestration of defensive iridoid glycosides (IGs), and viral infection load in field‐collected caterpillars using either P. lanceolata or a native plant, Chelone glabra (Plantaginaceae). We found that larvae using the exotic plant exhibited reduced immunocompetence, compositional differences in IG sequestration, and higher in situ viral burdens compared to those using the native plant. On both host plants, high IG sequestration was associated with reduced hemocyte concentration in the larval hemolymph, providing the first evidence of incompatibility between sequestered chemical defenses and the immune response (i.e., the "vulnerable host" hypothesis) from a field‐based study. However, despite this negative relationship between IG sequestration and cellular immunity, caterpillars with greater sequestration harbored lower viral loads. While survival of virus‐infected individuals decreased with increasing viral burden, it ultimately did not differ between the exotic and native plants. These results provide evidence that: (1) phytochemical sequestration may contribute to defense against pathogens even when immunity is compromised and (2) herbivore persistence on exotic plant species may be facilitated by sequestration and its role in defense against natural enemies. [ABSTRACT FROM AUTHOR]

Plant secondary (or specialized) metabolites mediate important interactions in both the rhizosphere and the phyllosphere. If and how such compartmentalized functions interact to determine plant-environment interactions is not well understood. Here, we investigated how the dual role of maize benzoxazinoids as leaf defenses and root siderophores shapes the interaction between maize and a major global insect pest, the fall armyworm. We find that benzoxazinoids suppress fall armyworm growth when plants are grown in soils with very low available iron but enhance growth in soils with higher available iron. Manipulation experiments confirm that benzoxazinoids suppress herbivore growth under iron-deficient conditions and in the presence of chelated iron but enhance herbivore growth in the presence of free iron in the growth medium. This reversal of the protective effect of benzoxazinoids is not associated with major changes in plant primary metabolism. Plant defense activation is modulated by the interplay between soil iron and benzoxazinoids but does not explain fall armyworm performance. Instead, increased iron supply to the fall armyworm by benzoxazinoids in the presence of free iron enhances larval performance. This work identifies soil chemistry as a decisive factor for the impact of plant secondary metabolites on herbivore growth. It also demonstrates how the multifunctionality of plant secondary metabolites drives interactions between abiotic and biotic factors, with potential consequences for plant resistance in variable environments. [ABSTRACT FROM AUTHOR]

The host acceptances of insects can be determined largely by detecting plant metabolites using insect taste. In the present study, we investigated the gustatory sensitivity and feeding behaviors of two closely related caterpillars, the generalist Helicoverpa armigera (Hübner) and the specialist H. assulta (Guenée), to different plant metabolites by using the single sensillum recording technique and the dual-choice assay, aiming to explore the contribution of plant metabolites to the difference of diet breadth between the two species. The results depicted that the feeding patterns of caterpillars for both plant primary and secondary metabolites were significantly different between the two Helicoverpa species. Fructose, glucose, and proline stimulated feedings of the specialist H. assulta , while glucose and proline had no significant effect on the generalist H. armigera. Gossypol and tomatine, the secondary metabolites of host plants of the generalist H. armigera , elicited appetitive feedings of this insect species but drove aversive feedings of H. assulta. Nicotine and capsaicin elicited appetitive feedings of H. assulta , but drove aversive feedings of H. armigera. For the response of gustatory receptor neurons (GRNs) in the maxillary styloconic sensilla of caterpillars, each of the investigated primary metabolites induced similar responding patterns between the two Helicoverpa species. However, four secondary metabolites elicited different responding patterns of GRNs in the two species, which is consistent with the difference of feeding preferences to these compounds. In summary, our results of caterpillars' performance to the plant metabolites could reflect the difference of diet breadth between the two Helicoverpa species. To our knowledge, this is the first report showing that plant secondary metabolites could drive appetitive feedings in a generalist insect species, which gives new insights of underscoring the adaptation mechanism of herbivores to host plants. [ABSTRACT FROM AUTHOR]

Context: The amount and composition of phytochemicals in forage plants influences habitat quality for wild herbivores. However, evaluating forage quality at fine resolutions across broad spatial extents (i.e., foodscapes) is challenging. Unmanned aerial systems (UAS) provide an avenue for bridging this gap in spatial scale. Objectives: We evaluated the potential for UAS technology to accurately predict nutritional quality of sagebrush (Artemisia spp.) across landscapes. We mapped seasonal forage quality across two sites in Idaho, USA, with different mixtures of species but similar structural morphotypes of sagebrush. Methods: We classified the sagebrush at both study sites using structural features of shrubs with object-based image analysis and machine learning and linked this classification to field measurements of phytochemicals to interpolate a foodscape for each phytochemical with regression kriging. We compared fine-scale landscape patterns of phytochemicals between sites and seasons. Results: Classification accuracy for morphotypes was high at both study sites (81–87%). Forage quality was highly variable both within and among sagebrush morphotypes. Coumarins were the most accurately mapped (r2 = 0.57–0.81), whereas monoterpenes were the most variable and least explained. Patches with higher crude protein were larger and more connected in summer than in winter. Conclusions: UAS allowed for a rapid collection of imagery for mapping foodscapes based on the phytochemical composition of sagebrush at fine scales but relatively broad extents. However, results suggest that a more advanced sensor (e.g., hyperspectral camera) is needed to map mixed species of sagebrush or to directly measure forage quality. [ABSTRACT FROM AUTHOR]

Plant secondary metabolites (PSMs) are a defense mechanism against herbivores, which in turn use detoxification metabolism to process ingested and absorbed PSMs. The feeding environment can cause changes in liver metabolism patterns and the gut microbiota. Here, we compared gut microbiota and liver metabolome to investigate the response mechanism of plateau zokors (Eospalax baileyi) to toxic plant Stellera chamaejasme (SC) in non-SC and SC grassland (-SCG and +SCG). Our results indicated that exposure to SC in the -SCG population increased liver inflammatory markers including prostaglandin (PG) in the Arachidonic acid pathway, while exposure to SC in the +SCG population exhibited a significant downregulation of PGs. Secondary bile acids were significantly downregulated in +SCG plateau zokors after SC treatment. Of note, the microbial taxa Veillonella in the -SCG group was significantly correlated with liver inflammation markers, while Clostridium innocum in the +SCG group had a significant positive correlation with secondary bile acids. The increase in bile acids and PGs can lead to liver inflammatory reactions, suggesting that +SCG plateau zokors may mitigate the toxicity of SC plants by reducing liver inflammatory markers including PGs and secondary bile acids, thereby avoiding liver damage. This provides new insight into mechanisms of toxicity by PSMs and counter-mechanisms for toxin tolerance by herbivores. [Display omitted] • Zokors inhabiting various grasslands exhibit distinct metabolic responses to toxic plants. • Zokors reduce chenodeoxycholic acid and prostaglandin to alleviate liver injury. • Veillonella associated with prostaglandin help zokors resist negative consequences of consuming toxic plants. • Liver-gut axis interaction was important for detoxify metabolism of toxic plants. [ABSTRACT FROM AUTHOR]

The koala is a specialist feeder with a diet consisting almost exclusively of potentially toxic eucalypt leaves. Monoterpenes, an abundant class of plant secondary metabolites in eucalypts, are highly lipophilic. Chronic absorption and systemic exposure can be anticipated for the koala, causing health effects in various ways when consumed in high amounts, but particularly causing alterations in immune function in this species. Therefore, careful leaf selection, efficient detoxification pathways, and other specialist adaptations are required to protect animals from acute intoxication. This is the first paper providing insight into the systemic exposure of koalas to these compounds. Profiles of six selected major monoterpenes were investigated in the ingesta of deceased koalas from four different regions of NSW and South-East Queensland. Concentrations of the same compounds were measured in lymphoid tissues of deceased koalas and in the blood of live koalas from other regions of NSW. Analytical methods included liquid extraction and solid-phase micro-extraction, followed by gas-chromatography/ mass-spectrometry. Concentrations in the ingesta of individual animals vary remarkably, though the average proportions of individual monoterpenes in the ingesta of animals from the four different regions are highly comparable. Blood concentrations of the selected monoterpenes also varied considerably. The highest blood concentrations were found for 1,8-cineole, up to 971 ng/ml. There was similarity between circulating monoterpene profiles and ingesta profiles. Based on the observed lack of similarity between blood and lymph tissue concentrations, individual monoterpenes either exhibit different affinities for lymphatic tissue compared to blood or their accumulation in blood and lymph tissue differs temporally. In general, blood monoterpene concentrations found in koalas were low compared to those reported in other marsupial eucalypt feeders, but significant concentrations of monoterpenes were detected in all samples analysed. This data on blood and lymphatic tissue monoterpene concentrations builds the fundamental groundwork for future research into the effects of dietary monoterpenes on various biological processes of specialist herbivores and into the significance of these animals' metabolic and behavioural strategies for coping with these compounds. We have shown that the systemic exposure of koalas to potentially anti-inflammatory eucalypt monoterpenes is continuous, and we provide data on physiological concentrations which will allow realistic future studies of the effects of monoterpenes on immune cell function. [ABSTRACT FROM AUTHOR]

Dietary specialists often reside in habitats that provide a high and predictable abundance of their primary food, which is usually difficult for other herbivores to consume because of high levels of plant toxins or structural impediments. Therefore, sympatric specialist and generalist herbivores may partition food resources within and among plants. We compared how a dietary specialist (pygmy rabbit, Brachylagus idahoensis) and generalist (mountain cottontail, Sylvilagus nuttallii) used sagebrush as a food resource during winter across 3 field sites in Idaho, United States, and in controlled feeding trials with captive rabbits. The proportion of sagebrush consumed by both rabbit species varied among sites, indicating that characteristics of sagebrush plants and the surrounding plant community influenced use of sagebrush. In addition, free-ranging and captive pygmy rabbits consumed a greater proportion of sagebrush and cropped smaller stem diameters with a greater proportion of sagebrush leaves (high monoterpenes, low fiber) relative to stems (low monoterpenes, high fiber) than did cottontails. Cottontails frequently discarded the leafy tips of sagebrush branches. Cottontails are more tolerant of fiber and less tolerant of sagebrush toxins than pygmy rabbits. Cottontails consumed large diameter stems, which diluted toxins in sagebrush but increased fiber intake and reduced digestible nitrogen intake. Pygmy rabbits are less tolerant of fiber but more able to detoxify and eliminate sagebrush toxins than cottontails. Pygmy rabbits consumed small diameter stems, which reduced fiber intake, but increased intake of toxins from sagebrush leaves. Although partitioning of stems and leaves within sagebrush plants may provide a mechanism for coexistence of specialist and generalist rabbits, higher-than-expected dietary overlap between both free-ranging and captive rabbits in winter might create resource competition in areas with high-density sympatric populations or low availability of sagebrush. In addition, these contrasting foraging strategies have the potential to influence dynamics of sagebrush communities over time. [ABSTRACT FROM AUTHOR]

Insect herbivores are important mediators of selection on traits that impact plant defense against herbivory and competitive ability. Although recent experiments demonstrate a central role for herbivory in driving rapid evolution of defense and competition-mediating traits, whether and how herbivory shapes heritable variation in these traits remains poorly understood. Here, we evaluate the structure and evolutionary stability of the G matrix for plant metabolites that are involved in defense and allelopathy in the tall goldenrod, Solidago altissima. We show that G has evolutionarily diverged between experimentally replicated populations that evolved in the presence versus the absence of ambient herbivory, providing direct evidence for the evolution of G by natural selection. Specifically, evolution in an herbivore-free habitat altered the orientation of G, revealing a negative genetic covariation between defense- and competition-related metabolites that is typically masked in herbivore-exposed populations. Our results may be explained by predictions of classical quantitative genetic theory, as well as the theory of acquisition-allocation trade-offs. The study provides compelling evidence that herbivory drives the evolution of plant genetic architecture. [ABSTRACT FROM AUTHOR]

Herbivores employ numerous strategies to reduce their exposure to toxic plant secondary chemicals (PSCs). However, the physiological mechanisms of PSC absorption have not been extensively explored. In particular, the absorption of PSCs via intestinal lymphatic absorption has been largely overlooked in herbivores, even though this pathway is well recognized for pharmaceutical uptake. Here, we investigated for the first time whether PSCs might be absorbed by lymphatic transport. We fed woodrats ( Neotoma albigula) diets with increasing concentrations of terpene-rich juniper ( Juniperus monosperma) either with or without a compound that blocks intestinal lymphatic absorption (Pluronic L-81). Woodrats consuming diets that contained the intestinal lymphatic absorption blocker exhibited increased food intakes and maintained higher body masses on juniper diets. Our study represents the first demonstration that PSCs may be absorbed by intestinal lymphatic absorption. This absorption pathway has numerous implications for the metabolism and distribution of PSCs in the systemic circulation, given that compounds absorbed via lymphatic transport bypass first-pass hepatic metabolism. The area of lymphatic transport of PSCs represents an understudied physiological pathway in plant-herbivore interactions. [ABSTRACT FROM AUTHOR]

The genus Brassica includes economically important oilseed and vegetable plants. A number of insect pests are known to infest these crops and cause significant losses in yield. The plants in the family Brassicaceae have multiple defense mechanisms to overcome or reduce the damage by these pests including defensive biochemicals. These Plant Secondary Metabolites (PSMs) involve myrosinase-glucosinolate system, different volatile compounds, lectins, phytoalexins and phytoanticipins. While some of the compounds are always present in the plant system, the others are synthesized after herbivore attack. These compounds can either directly protect the plant by having effect(s) on insects' biology/behaviour or indirectly by attracting the natural enemies of the pests. Because of these secondary compounds, Brassicas have the potential to be used in pest management such as biofumigation against soil pests, as trap crops and cover crops and hence, can be a part of push-pull strategy. An attempt has been made to review these compounds in Brassicas, their role in defense against insects and potential in pest management. [ABSTRACT FROM AUTHOR]

The vast number of plant secondary metabolites (PSMs) produced by higher plants has generated many efforts to exploit their potential for pest control. We performed a systematic literature search to retrieve relevant publications, and we evaluated these according to PSM groups to derive information about the potential for developing plant-derived rodent repellents. We screened a total of 54 publications where different compounds or plants were tested regarding rodent behavior/metabolism. In the search for widely applicable products, we recommend multi-species systematic screening of PSMs, especially from the essential oil and terpenoid group, as laboratory experiments have uniformly shown the strongest effects across species. Other groups of compounds might be more suitable for the management of species-specific or sex-specific issues, as the effects of some compounds on particular rodent target species or sex might not be present in non-target species or in both sexes. Although plant metabolites have potential as a tool for ecologically-based rodent management, this review demonstrates inconsistent success across laboratory, enclosure, and field studies, which ultimately has lead to a small number of currently registered PSM-based rodent repellents. [ABSTRACT FROM AUTHOR]

One function of the gut microbiota gaining recent attention, especially in herbivorous mammals and insects, is the metabolism of plant secondary metabolites (PSMs). We investigated whether this function exists within the gut communities of a specialist avian herbivore. We sequenced the cecal metagenome of the Greater Sage-grouse (Centrocercus urophasianus), which specializes on chemically defended sagebrush (Artemisia spp.). We predicted that the cecal metagenome of the sage-grouse would be enriched in genes associated with the metabolism of PSMs when compared to the metagenome of the domestic chicken. We found that representation of microbial genes associated with "Xenobiotic Degradation and Metabolism" was three-fold higher in the sage-grouse cecal metagenomes when compared to that of the domestic chicken. Further, we identified a complete metabolic pathway for the degradation of phenol to pyruvate, which was not detected in the metagenomes of the domestic chicken, bovine rumen, or 14 species of mammalian herbivores. Evidence of monoterpene degradation (a major class of PSMs in sagebrush) was less definitive, although we did detect genes for several enzymes associated with this process. Overall, our results suggest that the gut microbiota of specialist avian herbivores play a similar role to the microbiota of mammalian and insect herbivores in degrading PSMs. [ABSTRACT FROM AUTHOR]

Polyethylene glycol (PEG) has been used to study the intake and digestion of tannin-rich plants by mammalian herbivores because it preferentially binds to tannins. However, it is not clear whether the responses of herbivores to dietary PEG is due to increased protein availability from the release of tannin-bound protein, amelioration of tannin effects, or whether PEG also may bind to other compounds and change their activity in the gut. We used three native New Zealand tree species to measure the effect of PEG on the amount of foliage eaten by invasive common brushtail possums ( Trichosurus vulpecula) and on in vitro digestible nitrogen (available N). The addition of PEG increased the in vitro available N content of Weinmannia racemosa foliage, and possums ate significantly more PEG-treated foliage than untreated foliage. However, possums also ate more PEG-treated Fuchsia excorticata foliage, even though PEG did not increase in vitro available N in this species. Possums ate very little Melicytus ramiflorus, regardless of PEG treatment, even though M. ramiflorus contained the highest concentration of in vitro available N. These results prompted us to use PEG and a protein supplement, casein, to manipulate the available N concentration of diets containing ground eucalypt foliage, a well-studied food species for possums. Again, the response of possums to PEG was independent of changes in in vitro available N. In addition, altering the protein content of the diet via the addition of casein did not affect how much food the possums consumed. We conclude that the effects of PEG on dry matter intake by mammalian herbivores are not due solely to the release of tannin-bound protein. There is need for a better understanding of PEG-tannin interactions in order to ensure that the use of PEG in nutritional studies does not outstrip an understanding of its mechanisms of action. [ABSTRACT FROM AUTHOR]

Plants display extensive intraspecific variation in secondary metabolites. However, the selective forces shaping this diversity remain often unknown, especially below ground. Using Taraxacum officinale and its major native insect root herbivore Melolontha melolontha, we tested whether below-ground herbivores drive intraspecific variation in root secondary metabolites. We found that high M. melolontha infestation levels over recent decades are associated with high concentrations of major root latex secondary metabolites across 21 central European T. officinale field populations. By cultivating offspring of these populations, we show that both heritable variation and phenotypic plasticity contribute to the observed differences. Furthermore, we demonstrate that the production of the sesquiterpene lactone taraxinic acid β-D-glucopyranosyl ester (TA-G) is costly in the absence, but beneficial in the presence of M. melolontha, resulting in divergent selection of TA-G. Our results highlight the role of soil-dwelling insects for the evolution of plant defences in nature. [ABSTRACT FROM AUTHOR]

Identifying specific plant secondary metabolites that influence feeding behavior can be challenging, but a solid understanding of animal preferences can guide efforts. Common brushtail possums ( Trichosurus vulpecula) predominantly eat Eucalyptus species belonging to the subgenus Symphyomyrtus, and avoid eating those belonging to the Monocalyptus subgenus (also called subgenus Eucalyptus). Using an unbiased H NMR metabolomics approach, a previous study identified unsubstituted B ring flavanones in most species of monocalypts examined, whereas these compounds were absent from symphyomyrtles. We hypothesised that unsubstituted B ring flavanones act as feeding deterrents for common brushtail possums. In the current study, we tested this hypothesis by comparing how much possums ate of a basal diet, with diets containing one of four structurally related compounds; pinocembrin, flavanone (unsubstituted B ring flavanones), chrysin (the flavone analogue of pinocembrin), and naringenin (a flavanone with B ring substitution). We found that pinocembrin and flavanone deterred feeding relative to the basal diet, but that chrysin and naringenin did not at equivalent concentrations. Thus, unsubstituted B-ring flavanones may explain why brushtail possums avoid eating monocalypt species. Furthermore, small differences in the structure of secondary compounds can have a large impact on antifeedant properties. These results demonstrate that metabolomics can be a valuable tool for ecologists seeking to understand herbivore feeding preferences. [ABSTRACT FROM AUTHOR]

1. Although anthropogenic nitrogen (N) enrichment has significantly changed the growth, survival and reproduction of herbivorous insects, its effects on the defensive sequestration of secondary chemicals by insect herbivores are less well understood. Previous studies have shown that soil nutrient availability can affect sequestration directly through changing concentrations of plant defence chemicals, or indirectly through altering growth rates of herbivores. There has been less exploration of how nutrient deposition affects the consumption of secondary chemicals and subsequent sequestration efficiency. In the current study, the overall effect of soil N availability on cardenolide sequestration by the monarch caterpillar Danaus plexippus was examined. Specifically, the effects of soil nutrient availability on growth, consumption, excretion and sequestration efficiency of cardenolides by D. plexippus larvae fed on the tropical milkweed Asclepias curassavica were measured. 2. The results showed that soil N and phosphorus (P) fertilisation significantly reduced caterpillar growth rate and the sequestration efficiency of cardenolides by monarch caterpillars feeding on A. curassavica. The lowered sequestration efficiency was accompanied by higher concentrations of cardenolides in frass. Although the total cardenolide contents of caterpillars were lower under high N or P fertilisation levels, caterpillar cardenolide concentrations were constant across fertilisation treatments because of lower growth rates (and therefore lower body mass) under high fertilisation. It is concluded that anthropogenic N deposition may have multiple effects on insect herbivores, including their ability to defend themselves from predators with sequestered plant defences. [ABSTRACT FROM AUTHOR]

Abstract: We are using Flourensia cernua as a shrub model to study how terpenes affect livestock herbivory. Two experiments were conducted to examine distribution of volatile chemicals within a plant in an effort to minimize sample variability. In Experiment 1, leaves (current year''s growth) were collected from 20 tarbush plants. Two leaders were sampled from each of three positions (outer canopy, subcanopy, and basal) in all four quadrants (based on ordinal direction). In Experiment 2, 10 leaders of current year''s growth were removed from another 20 plants. Leaders were collected from the outer canopy of each quadrant and separated into thirds before removing leaves, thereby creating three leaf age categories. Volatile compounds were extracted with ethanol and analyzed with gas chromatography/mass spectrometry. Ninety-four chemicals (including 15 unknowns) were present on the leaf surface of F. cernua. Although 14 and 21 compounds differed (P < 0.05) among quadrants in Experiments 1 and 2, respectively, no consistent quadrant effect was detected in either study. Leaf position differed (P < 0.05) for 52 chemicals in Experiment 1 but outer canopy and subcanopy leaves differed for only 10 compounds. In Experiment 2, 63 compounds differed among leaf age categories. Immature leaves contained greater concentrations of 46 chemicals (P < 0.05) than intermediate or mature age categories, but intermediate and mature leaves differed for only seven compounds. Estimated total concentration (i.e., cumulative concentration of all compounds) was not affected by leaf position but varied among leaf age categories (P < 0.05; immature > intermediate > mature). Differences in leaf position were attributed about equally to mono- and sesquiterpenes, whereas leaf age differences were primarily due to sesquiterpenes. Leaf position and age both affect terpene concentration and sampling variability for tarbush. However, little difference was detected between subcanopy and canopy positions. Thus, by avoiding basal sprouts and sampling from the mid-point of current year''s growth, sampling variation should be minimal. [Copyright &y& Elsevier]

The co-evolved relationship between the chemical composition of plants and herbivory is fundamental in understanding diet selection of herbivores and their impacts on plants and ecosystems. However, the impact of plant secondary chemistry on mammalian herbivory is not fully understood. We investigated seasonal influences of phenolics with low molecular weight (e.g., flavonoids, salicylates) and plant morphology of the tea-leaved willow ( Salix phylicifolia L.) on moose ( Alces alces (L., 1758)) foraging. We analysed the relationship of different phenolic compounds in twigs and browsing in winter, and in leaves and the degree of leaf stripping in summer, and the role of plant morphology. In winter, higher concentrations of phenolics, e.g., myricetin-related compounds (belonging to the flavonoids), had a negative impact on herbivory by moose. This impact was not associated with a single compound but instead seemed to be more a result of synergistic or additive effects of different compounds. In contrast, the models for summer browsing showed a pronounced effect of plant morphology. Our analyses reveal differences in the relative importance of phenolics of low molecular weight and plant morphology on diet selection between seasons. These findings are relevant for understanding feeding decisions and mechanisms deterring mammalian herbivores known for shaping the vegetation on the ecosystem level. [ABSTRACT FROM AUTHOR]

Plants produce a variety of secondary metabolites (PSMs) that may be selective against herbivores. Yet, specialist herbivores may use PSMs as cues for host recognition, oviposition, and feeding stimulation, or for their own defense against parasites and predators. This summarizes a dual role of PSMs: deter generalists but attract specialists. It is not clear yet whether specialist herbivores are a selective force in the evolution of PSM diversity. A prerequisite for such a selective force would be that the preference and/or performance of specialists is influenced by PSMs. To investigate these questions, we conducted an oviposition experiment with cinnabar moths ( Tyria jacobaeae) and plants from an artificial hybrid family of Jacobaea vulgaris and Jacobaea aquatica. The cinnabar moth is a specialist herbivore of J. vulgaris and is adapted to pyrrolizidine alkaloids (PAs), defensive PSMs of these plants. The number of eggs and egg batches oviposited by the moths were dependent on plant genotype and positively correlated to concentrations of tertiary amines of jacobine-like PAs and some otosenine-like PAs. The other PAs did not correlate with oviposition preference. Results suggest that host plant PAs influence cinnabar moth oviposition preference, and that this insect is a potential selective factor against a high concentration of some individual PAs, especially those that are also involved in resistance against generalist herbivores. [ABSTRACT FROM AUTHOR]

Animal habitat selection is a process that functions at multiple, hierarchically structured spatial scales. Thus multi-scale analyses should be the basis for inferences about factors driving the habitat selection process. Vertebrate herbivores forage selectively on the basis of phytochemistry, but few studies have investigated the influence of selective foraging (i.e., fine-scale habitat selection) on habitat selection at larger scales. We tested the hypothesis that phytochemistry is integral to the habitat selection process for vertebrate herbivores. We predicted that habitats selected at three spatial scales would be characterized by higher nutrient concentrations and lower concentrations of plant secondary metabolites (PSMs) than unused habitats. We used the Greater Sage-Grouse (Centrocercus urophasianus), an avian herbivore with a seasonally specialized diet of sagebrush, to test our hypothesis. Sage-Grouse selected a habitat type (black sagebrush, Artemisia nova) with lower PSM concentrations than the alternative (Wyoming big sagebrush, A. tridentata wyomingensis). Within black sagebrush habitat, Sage-Grouse selected patches and individual plants within those patches that were higher in nutrient concentrations and lower in PSM concentrations than those not used. Our results provide the first evidence for multi-scale habitat selection by an avian herbivore on the basis of phytochemistry, and they suggest that phytochemistry may be a fundamental driver of habitat selection for vertebrate herbivores. [ABSTRACT FROM AUTHOR]

Generalist herbivores forage on a variety of plant species, allowing them to gain nutrients while limiting ingestion of harmful toxins. As the capacity to mix diets appears important for maximizing intake, the spatial scale in heterogeneity of food resources should influence the foraging behavior of herbivores. Our aim was to identify how the feeding strategy of a generalist mammalian herbivore, the common brushtail possum, responds to foods within a spatially defined environment. We evaluated foraging responses against increasing spatial separation between pairs of artificial diets that differed in flavor and toxin profile, to determine how distance and diet affect intake and behavior. Foraging responses were influenced by the type of diet or the degree of spatial separation between foods but not by their interaction. Diet influenced intake, time spent feeding, and feeding rate, but had no effect on nightly foraging interval, number of feeding bouts, or bout length. The number of switches between paired food resources and foraging efficiency (intake per unit distance, which accounts for the energetic costs of travelling), were influenced only by distance. Titrating foraging against a range of distances demonstrated how quickly foraging efficiency can decline in response to the spatial separation of food resources, highlighting the importance of spatial heterogeneity of plants within the home range of an herbivore. [ABSTRACT FROM AUTHOR]

We use modeling to determine the optimal relative plant carbon allocations between foliage, fine roots, anti-herbivore defense, and reproduction to maximize reproductive output. The model treats these plant components and the herbivore compartment as variables. Herbivory is assumed to be purely folivory. Key external factors include nutrient availability, degree of shading, and intensity of herbivory. Three alternative functional responses are used for herbivory, two of which are variations on donor-dependent herbivore (models 1a and 1b) and one of which is a Lotka-Volterra type of interaction (model 2). All three were modified to include the negative effect of chemical defenses on the herbivore. Analysis showed that, for all three models, two stable equilibria could occur, which differs from most common functional responses when no plant defense component is included. Optimal strategies of carbon allocation were defined as the maximum biomass of reproductive propagules produced per unit time, and found to vary with changes in external factors. Increased intensity of herbivory always led to an increase in the fractional allocation of carbon to defense. Decreases in available limiting nutrient generally led to increasing importance of defense. Decreases in available light had little effect on defense but led to increased allocation to foliage. Decreases in limiting nutrient and available light led to decreases in allocation to reproduction in models 1a and 1b but not model 2. Increases in allocation to plant defense were usually accompanied by shifts in carbon allocation away from fine roots, possibly because higher plant defense reduced the loss of nutrients to herbivory. [ABSTRACT FROM AUTHOR]

Abstract: Herbivores form preferences for substances that raise fitness such as nutritious foods and avoid those substances, which lower fitness such as plant secondary compounds (PSC). Nevertheless, some PSC at certain concentrations have the potential to raise fitness as they act against infective diseases such as parasitism. The objective of the present review is to assess whether parasitized farm animals are able to manifest self-medicative behaviors. The ability of herbivores to self-select these medicinal plant compounds emerges from the inherent adaptive nature of behavioral responses in living organisms. In order to manifest self-medication animals should experience discomfort caused by a certain illness and also relief associated with the mitigation of sickness brought about by the ingestion of a medicinal plant. Observational and controlled studies suggest such conditions, as well as self-medicative behaviors occur not only in wild but also in domestic herbivores. Self-medicative behaviors may allow individuals seek medicinal plant products when infected by parasites, even at times when the manager is not aware of the existence of parasitism in the flock. Self-medication may also allow for proper nutrition as animals offered choices will consume PSC-containing plants as well as nutritious and safe forages. Management programs should be geared at enhancing in herbivores the likelihood of the association medicine-relief from sickness and at favoring the transmission of self-medicative behavior across generations. [Copyright &y& Elsevier]

Abstract: Browsing by mammalian herbivores is a major problem in plantation forestry. Seedlings are most vulnerable to browsing during establishment, making protection crucial during this period. Aside from reducing herbivore numbers, browsing can be controlled through the application of tree guards or chemical repellents. These methods are generally short-term options. A promising, potentially longer-term method of reducing browsing damage on plantations is to plant individuals with enhanced natural browse resistance. This requires the development of a rapid, cost-effective means of identifying germplasm with enhanced resistance. Here we present such a screening methodology. We planted Eucalyptus globulus seedlings from 22 different seedlots in randomised blocks along edge rows of six operational plantations. Seedlings were monitored for mammal browsing damage and growth for 2years. Natural variation between E. globulus seedlots resulted in significant variation in the extent to which they were browsed. Differential browsing was evident just 10weeks after planting, and caused a significant effect on tree growth after 2years. Differential browsing was thought to be due to variation in levels of plant secondary metabolites (PSMs). The identification and selective planting of high PSM seedlots in high risk areas could be an effective management tool to reduce browsing in plantation forestry. [Copyright &y& Elsevier]

Abstract: Rangelands throughout the world contain varying but often substantial proportions of shrubs. Shrubs are generally heavily chemically defended, and herbivores must either contend with their plant secondary metabolites (PSM) or avoid a significant component of the available forage. Browsing ruminants are exposed to thousands of chemicals in infinite combinations and concentrations that are constantly changing both temporally and spatially. The success with which a herbivore navigates this complex environment is in part attributed to its ability to cope with PSM. Plant secondary metabolites can affect a number of physiological and metabolic processes (e.g., altered microbial activity, reduced digestion, compromised acid/base balance, toxicity), although negative consequences to the herbivore range from harmless to lethal, depending factors such as dose, animal species, plane of nutrition, and physiological state. Herbivores have a variety of intertwined mechanisms to cope with consumption of PSM, ranging from physiological (e.g., salivary proteins, detoxification pathways) to behavioral (e.g., avoidance, regulation of intake below critical threshold, cautious sampling, altering size and pattern of feeding bouts, diet switching, consuming diverse and/or complementary diets). Secondary compounds may affect requirements for nutrients (e.g., protein, minerals, and glucose) and water, and may alter basal metabolic rate. Energy requirements may also increase to accommodate increased travel to water and supplementation sites to counter these negative effects, particularly on arid rangelands. A number of management strategies exist for minimizing the negative effects of PSM consumption on livestock. Supplementation to replace nutrients depleted during detoxification (e.g., amino acids and glucose), additives such as PEG and charcoal to reduce absorption and increase excretion of PSM, maintaining animals in good body condition, and behavioral modifications (e.g., diet training) are among the potential management options for enhancing the ability of ruminants to cope with PSM when browsing shrubby ecosystems. [Copyright &y& Elsevier]

Abstract: Plants provide herbivores with an array of chemicals with the potential to improve health and well-being. For instance, phytochemicals, known as secondary compounds, which protect plants from consumers and pests can adversely affect cellular and metabolic processes in herbivores, but at low doses and in appropriate mixtures, they can have beneficial effects on animal nutrition and health, though the latter has not been explored in great breadth or depth. In this review, we summarize the potential impact of natural plant products on immunomodulation and other therapeutic effects in herbivores. Development of preventative strategies to help animals resist disease would be a more economical, ecological and socially effective long-term healthcare strategy than treating diseases. In this realm, immunomodulation promoted by forages emerges as an interesting alternative and complement to chemotherapy. The challenge for feeding systems will be to incorporate mixes of plants with bioactive properties in ways that enhance health without compromising animal production and well-being. A solution to this challenge may involve developing management programs that acknowledge the ability of animals to learn about the beneficial effects of diverse foods. [Copyright &y& Elsevier]

• Formylated phloroglucinols (FPCs) are key defensive compounds that influence herbivory by mammals and arthropods in eucalypts. However, the genetic architecture underlying variation in their levels remains poorly understood. • Quantitative trait loci (QTL) analysis for the concentrations of two major FPCs, sideroxylonal A and macrocarpal G, was conducted using juvenile leaves from 112 clonally duplicated progenies from an outcross F2 of Eucalyptus globulus. • Two unlinked QTL were located for macrocarpal, while another unlinked QTL was located for sideroxylonal. The sideroxylonal QTL collocated with one for total sideroxylonal previously reported using adult Eucalyptus nitens foliage, providing independent validation in a different evolutionary lineage and a different ontogenetic stage. • Given the potential widespread occurrence of these QTL, their ontogenetic stability, and their impact on a range of dependent herbivores, it is possible that they have extended phenotypic effects in the Australian forest landscape. [ABSTRACT FROM AUTHOR]

1. Comparisons of adult and juvenile plant stages are often confounded by other factors such as differences in height between plants. However, these factors can be teased apart using common garden experiments and appropriate sampling designs. 2. Using paired sampling of upper and lower canopies of Eucalyptus nitens trees, which were growing in a common environment trial and only one of which had undergone the transition to adult foliage, this study assessed the effects of ontogeny and canopy position on the feeding preferences of Common Brushtail Possums ( Trichosurus vulpecula) in captive feeding trials under choice conditions. 3. Possums preferred juvenile foliage over adult foliage. In contrast, canopy position did not influence preferences. 4 Adult foliage contained significantly less sideroxylonals (a compound known to deter herbivory by Brushtail Possums), but was significantly tougher (with thicker cuticle, more fibre, more lignin and a higher percentage dry mass) than juvenile foliage, suggesting that leaf toughness was more significant than defensive chemistry in influencing intake by possums. Adult foliage did not differ significantly from juvenile foliage in nitrogen, total phenolics or essential oil concentration. 5. Greater resistance to herbivory of adult foliage compared with juvenile foliage by a significant browser of eucalypt tree foliage could contribute to selection for an earlier change from juvenile to adult foliage in areas that are heavily browsed by possums. [ABSTRACT FROM AUTHOR]

Plant secondary metabolites (PSMs) offer plants chemical defences against herbivores, and are known to influence intake and diet choice in both insect and mammalian herbivores. However, there is limited knowledge regarding how PSMs influence herbivore foraging decisions. Herbivore foraging decisions, in turn, directly impact on which individual plants, and plant species, are selected for consumption. We took advantage of the natural variation in sideroxylonal concentrations in the foliage of Eucalyptus melliodora (Cunn. ex Schauer) to investigate feeding patterns of a marsupial folivore, the common ringtail possum, Pseudocheirus peregrinus (Boddaert 1785). Foliage, collected from six trees, contained between 0.32 and 12.97 mg g-DM-1 sideroxylonal. With increasing sideroxylonal concentrations, possums decreased their total intake, rate of intake and intake per feeding bout, and increased their cumulative time spent feeding. Possums did not alter their total feeding time, number of feeding bouts or time per feeding bout in response to increasing sideroxylonal concentrations. Results demonstrate important behavioural changes in foraging patterns in response to sideroxylonal. These behavioural changes have important implications, in relation to altered foraging efficiency and potential predation risk, for herbivores foraging in the field. As a result, the spatial distribution of dietary PSMs across a landscape may directly influence herbivore fitness, and ultimately habitat selection of mammalian herbivores. [ABSTRACT FROM AUTHOR]

The diamondback moth (DBM), Plutella xylostella (Lepidoptera: Plutellidae) is a very destructive crucifer-specialized pest that has resulted in significant crop losses worldwide. DBM is well attracted to glucosinolates (which act as fingerprints and essential for herbivores in host plant recognition) containing crucifers such as wintercress, Barbarea vulgaris (Brassicaceae) despite poor larval survival on it due to high-to-low concentration of saponins and generally to other plants in the genus Barbarea. B. vulgaris build up resistance against DBM and other herbivorous insects using glucosinulates which are used in plant defense. Aside glucosinolates, Barbarea genus also contains triterpenoid saponins, which are toxic to insects and act as feeding deterrents for plant specialist herbivores (such as DBM). Previous studies have found interesting relationship between the host plant and secondary metabolite contents, which indicate that attraction or resistance to specialist herbivore DBM, is due to higher concentrations of glucosinolates and saponins in younger leaves in contrast to the older leaves of Barbarea genus. As a response to this phenomenon, herbivores as DBM has developed a strategy of defense against these plant biochemicals. Because there is a lack of full knowledge in understanding bioactive molecules (such as saponins) role in plant defense against plant herbivores. Thus, in this review, we discuss the role of secondary plant metabolites in plant defense mechanisms against the specialist herbivores. In the future, trials by plant breeders could aim at transferring these bioactive molecules against herbivore to cash crops. [ABSTRACT FROM AUTHOR]