Parasites as ecosystem modulators: foliar pathogens suppress top-down effects of large herbivores.

Parasites can catalyze or inhibit interactions between their hosts and other species, but the ecosystem-level effects of such interaction modifications are poorly understood. We conducted a large-scale field experiment in temperate grasslands of China to understand how foliar fungal pathogens influenced top-down effects of cattle on plant diversity and productivity. When foliar pathogens were suppressed, cattle grazing strongly reduced biomass of the dominant grass, Leymus chinensis, generating competitive release that significantly increased community-level species richness and evenness. In the absence of grazing, pathogen attack on L. chinensis had no measurable effect on host biomass. However, pathogens disrupted top-down effects of herbivory by inhibiting grazing effects on plant biomass and species richness. Mechanistically, fungal pathogens were linked to increased alkaloid and reduced nitrogen levels in leaf tissue, which appeared to deter cattle grazing on L. chinensis. In conclusion, foliar pathogens can suppress top-down effects of large herbivores on grassland community composition and ecosystem function by modifying the strength of their host's interactions with dominant consumers. Parasites may act as modulators of ecosystem function when their direct effects on host abundance are overshadowed by powerful influences on host traits that modify their interactions with competitors, herbivores, or predators.

The role of herbivory in the macroevolution of vertebrate hormone dynamics.

Vertebrates have high species-level variation in circulating hormone concentrations, and the functional significance of this variation is largely unknown. We tested the hypothesis that interspecific differences in hormone concentrations are partially driven by plant consumption, based on the prediction that herbivores should have higher basal hormone levels to 'outcompete' plant endocrine disruptors. We compared levels of glucocorticoids (GCs), the hormones with the most available data, across 166 species. Using phylogenetically informed comparisons, we found that herbivores had higher GC levels than carnivores. Furthermore, we found that the previously described negative relationship between GC levels and body mass only held in herbivores, not carnivores, and that the effect of diet was greatest at extreme body sizes. These findings demonstrate the far-reaching effects of diet on animal physiology, and provide evidence that herbivory influences circulating hormone concentrations. We urge future direct testing of the relationship between phytochemical load and GC levels.

Digesta passage in nondomestic ruminants: Separation mechanisms in 'moose-type' and 'cattle-type' species, and seemingly atypical browsers.

Ruminants have been classified as having a 'moose-type' or 'cattle-type' digestive physiology. 'Cattle-type' ruminants have a clear difference in the mean retention time (MRT) of fluid vs. small particles in the reticulorumen (RR), with a high 'selectivity factor' (SF = MRTparticle/ MRTfluid, >1.80), and are typically grazers and intermediate feeders. 'Moose-type' ruminants have lower SF (<1.80), possibly because of defensive salivary proteins that constrain amounts of (high-viscosity) saliva, and are typically restricted to browsing. To further contribute to testing this physiology-diet correlation, we performed 55 individual passage measurements in 4/6 species that have/have not been investigated previously, respectively. Co-EDTA was used as a solute (fluid) and Cr-mordanted hay particles (<2 mm) as particle markers. Results are related to the percentage of grass in the natural diet taken from the literature. Moose (Alces alces, n = 4 on 4 to 5 diets each and n = 2 on a single diet, 5% grass, SF 1.46 ±â€¯0.22) and giraffe (Giraffa camelopardalis, n = 3 on 3 to 5 diets each, 1%, 1.42 ±â€¯0.23) as classical 'moose-type', and cattle (Bos taurus, n = 2, 70%, 2.04) as classical 'cattle-type' ruminants yielded results similar to those previously published, as did waterbuck (Kobus ellipsiprymnus, n = 5, 84%, 2.46 ±â€¯0.49), corroborating that the SF represents, to a large extent, a species-specific characteristic. Results in oryx (Oryx leucoryx, n = 1, 75%, 2.60) and sitatunga (Tragelaphus spekii, n = 4, 68%, 1.81 ±â€¯0.21) correspond to the concept of 'cattle-type' ruminants being grazers or intermediate feeders. However, European bison (Bison bonasus, n = 1, 10%, 2.74), nyala (T. angasii, n = 6, 20%, 1.95 ±â€¯0.25), bongo (T. eurycerus, n = 3, 13%, 2.39 ±â€¯0.54) and gerenuk (Litocranius walleri, n = 1, 0%, 2.25) appear as 'cattle-type' ruminants, yet have a browse-dominated diet. While the results do not challenge the view that a 'moose-type' digestive physiology is an adaptation to browse diets, they indicate that it may not be the only adaptation that enables ruminants to use browse. Apparently, a 'cattle-type' digestive physiology with a high SF does not necessarily preclude a browsing diet niche. High-SF browsers might have the benefit of an increased harvest of RR microbiota and grit removal prior to rumination; how they defend themselves against secondary plant compounds in browse remains to be investigated.

Isoxanthohumol, a constituent of hop (Humulus lupulus L.), increases stress resistance in Caenorhabditis elegans dependent on the transcription factor DAF-16.

PURPOSE: The flavanone isoxanthohumol (IX) has gained attention as antioxidative and chemopreventive agent, but the molecular mechanism of action remains unclear. We investigated effects of this secondary plant compound in vivo using the model organism Caenorhabditis elegans. METHODS: Adult C. elegans nematodes were incubated with IX, and then, the stress resistance was analysed in the SYTOX assay; lifespan was monitored by touch-provoked movement method, the amount of reactive oxygen species (ROS) was measured in the DCF assay, and the nuclear localisation of the transcription factor DAF-16 was analysed by using a transgenic strain. By the use of a DAF-16 loss-of-function strain, we analysed whether the effects are dependent on DAF-16. RESULTS: IX increases the resistance of the nematode against thermal stress. Additionally, a reduction in ROS in vivo was caused by IX. Since the flavanone only has a marginal radical-scavenging capacity (TEAC assay), we suggest that IX mediates its antioxidative effects indirectly via activation of DAF-16 (homologue to mammalian FOXO proteins). The nuclear translocation of this transcription factor is increased by IX. In the DAF-16-mutated strain, the IX-mediated increase in stress resistance was completely abolished; furthermore, an increased formation of ROS and a reduced lifespan was mediated by IX. CONCLUSION: IX or a bacterial metabolite of IX causes antioxidative effects as well as an increased stress resistance in C. elegans via activation of DAF-16. The homologous pathway may have implications in the molecular mechanism of IX in mammals.

On the study of plant defence and herbivory using comparative approaches: how important are secondary plant compounds.

Species comparisons are a cornerstone of biology and there is a long tradition of using the comparative framework to study the ecology and evolution of plant defensive traits. Early comparative studies led to the hypothesis that plant chemistry plays a central role in plant defence, and the evolution of plant secondary chemistry in response to insect herbivory remains a classic example of coevolution. However, recent comparative work has disagreed with this paradigm, reporting little connection between plant secondary chemicals and herbivory across distantly related plant taxa. One conclusion of this new work is that the importance of secondary chemistry in plant defence may have been generally overstated in earlier research. Here, we attempt to reconcile these contradicting viewpoints on the role of plant chemistry in defence by critically evaluating the use and interpretation of species correlations as a means to study defence-herbivory relationships. We conclude that the notion that plant primary metabolites (e.g. leaf nitrogen content) are the principal determinants of herbivory (or the target of natural selection by herbivores) is not likely to be correct. Despite the inference of recent community-wide studies of herbivory, strong evidence remains for a prime role of secondary compounds in plant defence against herbivores.

The Lignan Pinoresinol Induces Nuclear Translocation of DAF-16 in Caenorhabditis elegans but has No Effect on Life Span.

The lignan pinoresinol is a constituent of flaxseed, sesame seeds and olive oil. Because of different molecular effects reported for this compound, e.g. antioxidative activity, pinoresinol is suggested to cause positive effects on humans. Because experimental data are limited, we have analysed the effects of the lignan on the nematode Caenorhabditis elegans: in spite of a strong antioxidative capacity detected in an in vitro assay, no antioxidative effects were detectable in vivo. In analogy to this result, no modulation of the sensitivity against thermal stress was detectable. However, incubation with pinoresinol caused an enhanced nuclear accumulation of the transcription factor DAF-16 (insulin/IGF-like signalling pathway). Using a strain with an enhanced oxidative stress level (mev-1 mutant), we clearly see an increase in stress resistance caused by this lignan, but no change in reactive oxygen species. Furthermore, we investigated the effects of pinoresinol on the life span of the nematode, but no modulation was found, neither in wild-type nor in mev-1 mutant nematodes. These results suggest that pinoresinol may exert pharmacologically interesting effects via modulation of the insulin-like signalling pathway in C. elegans as well as in other species like mammals due to the evolutionary conservation of this signalling pathway.

Xenobiotic metabolism of plant secondary compounds in the English grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae).

Plant secondary compounds have been documented to be deleterious to insects and other herbivores in diverse ways. In this study, the effect of catechol (phenolics), gramine (alkaloid) and L-ornithine-HCI (non-protein amino acid) on the activities of xenobiotic metabolizing enzymes in English grain aphid, Sitobion avenae, was evaluated. Phase I enzymes investigated in this study included carboxylesterase (CarE), and oxidoreductase, whereas Phase II enzymes were represented by glutathione S-transferase (GST). In general, CarE and GST activities in S. avenae were positively correlated with the concentration of plant secondary compounds in artificial diets. Oxidoreductase activity, however, displayed a different profile. Specifically, peroxidase (POD) and polyphenol oxidase (PPO) activities in S. avenae were positively correlated with concentrations of dietary catechol and gramine, respectively, whereas catalase (CAT) activity was significantly suppressed by the higher concentration of catechol, gramine and L-ornithine-HCl. These combined results suggest that CarE and GST in S. avenae are key enzymes to breakdown a broad spectrum of plant secondary compounds, whereas oxidoreductase, including PPO and POD, degrades specific groups of plant secondary compounds.

Dietary Challenges for Parasitoid Wasps (Hymenoptera: Ichneumonoidea); Coping with Toxic Hosts, or Not?

Many insects defend themselves against predation by being distasteful or toxic. The chemicals involved may be sequestered from their diet or synthesized de novo in the insects' body tissues. Parasitoid wasps are a diverse group of insects that play a critical role in regulating their host insect populations such as lepidopteran caterpillars. The successful parasitization of caterpillars by parasitoid wasps is contingent upon their aptitude for locating and selecting suitable hosts, thereby determining their efficacy in parasitism. However, some hosts can be toxic to parasitoid wasps, which can pose challenges to their survival and reproduction. Caterpillars employ a varied array of defensive mechanisms to safeguard themselves against natural predators, particularly parasitoid wasps. These defenses are deployed pre-emptively, concurrently, or subsequently during encounters with such natural enemies. Caterpillars utilize a range of strategies to evade detection or deter and evade attackers. These tactics encompass both measures to prevent being noticed and mechanisms aimed at repelling or eluding potential threats. Post-attack strategies aim to eliminate or incapacitate the eggs or larvae of parasitoids. In this review, we investigate the dietary challenges faced by parasitoid wasps when encountering toxic hosts. We first summarize the known mechanisms through which insect hosts can be toxic to parasitoids and which protect caterpillars from parasitization. We then discuss the dietary adaptations and physiological mechanisms that parasitoid wasps have evolved to overcome these challenges, such as changes in feeding behavior, detoxification enzymes, and immune responses. We present new analyses of all published parasitoid-host records for the Ichneumonoidea that attack Lepidoptera caterpillars and show that classically toxic host groups are indeed hosts to significantly fewer species of parasitoid than most other lepidopteran groups.

Micronutrients and Markers of Oxidative Stress and Inflammation Related to Cardiometabolic Health: Results from the EHES-LUX Study.

Metabolic syndrome (MetS) characteristics include chronic inflammation and elevated oxidative stress. This study assessed associations between circulating concentrations of micronutrients/phytochemicals and inflammatory/oxidative stress markers with MetS and MetS components. Adults (N = 606) from the European Health Examination Survey in Luxembourg (2013-2015) were randomly selected. We performed a multivariable logistic regression model using the least absolute shrinkage and selection operator to identify MetS-associated variables. Participants with MetS had higher concentrations of C-reactive protein (CRP), 8-iso-prostaglandin F2α, leptin, insulin, and vitamins E/A, but lower concentrations of adiponectin, beta-carotene, and oxidized low-density lipoprotein. A one-unit increase in log-CRP was associated with 51% greater odds of MetS (OR = 1.51 (95% CI: 1.16, 1.98)). Adults with a one-unit increase in log-leptin were 3.1 times more likely to have MetS (3.10 (2.10, 4.72)). Women with a one-unit increase in vitamin A were associated with 3% increased odds of MetS (1.03 (1.01, 1.05)), while those with a one-unit increase in log-adiponectin were associated with 82% decreased odds (0.18 (0.07, 0.46)). Chronic inflammation best characterized adults with MetS, as CRP, adiponectin, and leptin were selected as the main MetS determinants. Micronutrients did not seem to affect MetS, except for vitamin A in women.

Dietary ursolic acid improves health span and life span in male Drosophila melanogaster.

The health and life span of Drosophila melanogaster are partly determined by intestinal barrier integrity, metabolic rate as well as stress response and the expression of longevity-associated genes, depending on genetic and dietary factors. Ursolic acid (UA) is a naturally occurring triterpenoid exhibiting potential antimicrobial, anti-inflammatory, and antiobesity activity and counteracting age-related deficits in muscle strength. In this study, UA was dietarily administered to w1118 D. melanogaster which significantly elongated the health and life span of males. Spargel (srl) is the Drosophila orthologue of mammalian peroxisome proliferator-activated receptor-gamma coactivator 1 α(PGC1α), an important regulator of energy homeostasis and mitochondrial function. Our results indicate that the health-promoting effect of UA, demonstrated by a significant increase in climbing activity, occurs via an upregulation of srl expression leading to a metabolic shift in the fly without reducing fecundity or gut integrity. Moreover, UA affected the flies' microbiota in a manner that contributed to life span extension. Srl expression and microbiota both seem to be affected by UA, as we determined by using srl-mutant and axenic flies. © 2018 BioFactors, 45(2):169-186, 2019.

Sea Buckthorn Oil as a Valuable Source of Bioaccessible Xanthophylls.

Sea buckthorn oil, derived from the fruits of the shrub, also termed seaberry or sandthorn, is without doubt a strikingly rich source of carotenoids, in particular zeaxanthin and ß-carotene. In the present study, sea buckthorn oil and an oil-in-water emulsion were subjected to a simulated gastro-intestinal in vitro digestion, with the main focus on xanthophyll bioaccessibility. Zeaxanthin mono- and di-esters were the predominant carotenoids in sea buckthorn oil, with zeaxanthin dipalmitate as the major compound (38.0%). A typical fatty acid profile was found, with palmitic (49.4%), palmitoleic (28.0%), and oleic (11.7%) acids as the dominant fatty acids. Taking into account the high amount of carotenoid esters present in sea buckthorn oil, the use of cholesterol esterase was included in the in vitro digestion protocol. Total carotenoid bioaccessibility was higher for the oil-in-water emulsion (22.5%) compared to sea buckthorn oil (18.0%) and even higher upon the addition of cholesterol esterase (28.0% and 21.2%, respectively). In the case of sea buckthorn oil, of all the free carotenoids, zeaxanthin had the highest bioaccessibility (61.5%), followed by lutein (48.9%), making sea buckthorn oil a potential attractive source of bioaccessible xanthophylls.

The resveratrol derivatives trans-3,5-dimethoxy-4-fluoro-4'-hydroxystilbene and trans-2,4',5-trihydroxystilbene decrease oxidative stress and prolong lifespan in Caenorhabditis elegans.

OBJECTIVES: Resveratrol (trans-3,4',5-trihydroxystilbene (1)) was previously shown to extend the lifespan of different model organisms. However, its pharmacological efficiency is controversially discussed. Therefore, the bioactivity of four newly synthesized stilbenes (trans-3,5-dimethoxy-4-fluoro-4'-hydroxystilbene (3), trans-4'-hydroxy-3,4,5-trifluorostilbene (4), trans-2,5-dimethoxy-4'-hydroxystilbene (5), trans-2,4',5-trihydroxystilbene (6)) was compared to (1) and pterostilbene (trans-3,5-dimethoxy-4'-hydroxystilbene (2)) in the established model organism Caenorhabditis elegans. METHODS: Trolox equivalent antioxidant capacity (TEAC), 2',7'-dichlorofluorescein (DCF), thermotolerance assays, C. elegans lifespan analyses. KEY FINDINGS: All compounds exert a strong in-vitro radical scavenging activity (6 > 1 > 5 > 2 = 3 = 4), but in vivo, only (3) and (6) reduce reactive oxygen species (ROS) accumulation. Furthermore, (3) and (6) increased the mobility of aged nematodes and prolonged their mean lifespans, while these compounds decreased the thermal stress resistance. Using daf-16 (FoxO), skn-1 (Nrf2) and sir-2.1 (sirtuin) loss-of-function mutant strains, the in vivo antioxidant effects of compounds (3) and (6) were abolished, showing the necessity of these evolutionary highly conserved factors. However, short-time treatment with stilbenes (3) and (6) did not modulate the cellular localization of the transcription factors DAF-16 and SKN-1. CONCLUSION: In contrast to resveratrol, the synthetic stilbene derivatives (3) and (6) increase the lifespan of C. elegans, rendering them promising candidates for pharmacological anti-ageing purposes.

Baicalein modulates stress-resistance and life span in C. elegans via SKN-1 but not DAF-16.

The flavonoid baicalein has been demonstrated to be an activator of the transcription factor Nrf2 in mammalian cell lines. We show that it further modulates the Nrf2 homolog SKN-1 in Caenorhabditis elegans and by this pathway mediates beneficial effects in the nematode: baicalein enhances the resistance of C. elegans against lethal thermal and sodium arsenite stress and dose-dependently prolongs the life span of the nematode. Using RNA interference against SKN-1 we were able to show that the induction of longevity and the enhanced stress-resistance were dependent on this transcription factor. DAF-16 (homolog to mammalian FOXO) is another pivotal aging-related transcription factor in the nematode. We demonstrate that DAF-16 does not participate in the beneficial effects of baicalein: since baicalein causes no increase in the nuclear translocation of DAF-16 (DAF-16::GFP expressing strain, incubation time: 1h) and it still induces longevity even in a DAF-16 loss-of-function strain, we conclude, that baicalein increases stress-resistance and life span in C. elegans via SKN-1 but not DAF-16.

Determinants of seasonal feeding of the generalist snail Arianta arbustorum at six sites dominated by Senecioneae.

We investigated the diet constituents of Arianta arbustorum by means of faecal analysis and regressed their quantity on the availability and quality of food plants. We studied six sites, all dominated by plants from the Asteraceae tribe Senecioneae, early and later in the growing season, predicting that the snail's food choice would be determined by the presence of the different secondary compounds found in the Senecioneae. The snails consumed less green plant material and more leaf litter in May than in July. Plant water and nitrogen content were higher in May. For the May samples, regressions were only significant at sites that excluded Adenostyles alliariae and Adenostyles alpina; in July, the regression fit was higher at all sites. Of the measured plant parameters, only quantity (availability) was a consistently significant variable in the regressions. Circumstantial evidence suggested that plant secondary compounds had a major influence on snail food choice: fresh-plant consumption increased over the season, as the concentration of many secondary compounds decreased; both plant availability and quality could only explain about half of the observed variation in snail feeding, which argues for other hidden factors influencing snail feeding; the dynamics of feeding of the various senecionean plants were such that A. alliariae and Senecio sp. had a seasonal acceptance whereas Petasites albus was always and A. alpina never accepted, and, finally, the detection of a number of snail-deterrent fractions in A. alpina leaves, a plant which was never found to be consumed in this study.

Behavioural responses of Frankliniella occidentalis Pergande larvae to methyl jasmonate and cis-jasmone.

The larval stages of Frankliniella occidentalis Pergande (Thysanoptera: Thripidae) cause more direct feeding damage to plants than the adults. We, therefore, investigated the behaviour-modifying effects on second instar larvae of two jasmonic acid derivatives. The artificial application of methyl jasmonate and cis-jasmone, both at 1 % concentration, deterred the larvae from settling in a dual choice bean leaf disc assay. We observed a dose-dependent feeding deterrence of both jasmonates and calculated the concentration required to reduce the feeding damage by 50 % relative to the control treatment (FDC50) for each jasmonate. The feeding damage was reduced by the application of cis-jasmone at 1 % concentration, but not by the jasmonates at the respective FDC50 in no-choice leaf disc bioassays. However, significantly more larvae left jasmonate-treated whole potted bean plants by migrating to the soil compared with control plants. Our results may be exploited extending behavioural manipulation by using plant compounds in thrips control programmes to the full lifecycle of the pest. Plant compounds could be used in integrated and biological pest management strategies against F. occidentalis in combination with the application of various above and below ground control measures.

Preference of Neoseiulus californicus (Acari: Phytoseiidae) for volatiles of Bt maize induced by multiple herbivory

ABSTRACT Plant indirect induced defenses against herbivores are characterized by the production of plant volatiles that to attract natural enemies. The objective of this study was to evaluate whether the attack of the two-spotted spider mite Tetranychus urticae or the multiple herbivory of T. urticae together with the fall armyworm Spodoptera frugiperda are able to elicit indirect induced defense in conventional and Bt maize plants. The experiment was carried out in the laboratory using Y-tube olfactometer, evaluating the predatory mite Neoseiulus californicus olfactory preference for plant volatiles. The treatments involved: Clean Conventional Plant; Conventional Plant Infested with T. urticae; Clean Bt Plant; Bt Plant Infested with T. urticae; Conventional Plant Infested with T. urticae + fall armyworm; Bt Plant Infested with T. urticae + fall armyworm. For the chemical analyzes the Trace GC Ultra gas chromatograph was used, paired with the Polaris Q mass spectrometer, GC - MS system. Neoseiulus californicus was also unable to distinguish between volatiles from both conventional and Bt infested maize plants. Moreover, there was no discrimination of the predator mite between plants under single and multiple infestations, both in conventional and Bt maize. When comparing conventional and Bt plants, both with multiple infestation, the predator mite N. californicus had no preference among these sources of odors. However, there was observed chemical changes of the volatiles among the groups of plants studied. Thus, it is suggested that the groups of plants under study have chemical modifications, but they are not able to attract N. californicus. In addition, Bt plants response was similar to conventional plants on attracting N. californicus.

Plant systemic induced responses mediate interactions between root parasitic nematodes and aboveground herbivorous insects.

Insects and nematodes are the most diverse and abundant groups of multicellular animals feeding on plants on either side of the soil-air interface. Several herbivore-induced responses are systemic, and hence can influence the preference and performance of organisms in other plant organs. Recent studies show that plants mediate interactions between belowground plant parasitic nematodes (PPNs) and aboveground herbivorous insects. Based on the knowledge of plant responses to pathogens, we review the emerging insights on plant systemic responses against root-feeding nematodes and shoot-feeding insects. We discuss the potential mechanisms of plant-mediated indirect interactions between both groups of organisms and point to gaps in our knowledge. Root-feeding nematodes can positively or negatively affect shoot herbivorous insects, and vice versa. The outcomes of the interactions between these spatially separated herbivore communities appear to be influenced by the feeding strategy of the nematodes and the insects, as well as by host plant susceptibility to both herbivores. The potential mechanisms for these interactions include systemic induced plant defense, interference with the translocation and dynamics of locally induced secondary metabolites, and reallocation of plant nutritional reserves. During evolution, PPNs as well as herbivorous insects have acquired effectors that modify plant defense responses and resource allocation patterns to their advantage. However, it is also known that plants under herbivore attack change the allocation of their resources, e.g., for compensatory growth responses, which may affect the performance of other organisms feeding on the plant. Studying the chemical and molecular basis of these interactions will reveal the molecular mechanisms that are involved. Moreover, it will lead to a better understanding of the ecological relevance of aboveground-belowground interactions, as well as support the development of sustainable pest management technologies.