Rutin: a phagostimulant for the polyphagous acridid Schistocerca americana

Rutin was demonstrated to be a phagostimulant for the grasshopper Schistocerca americana across a very wide concentration range. The effect was not maintained over a period of days if the insects had already ingested a large amount, but when individuals were given different concentrations on disks as supplements to their lettuce diet, they tended to self select a moderate quantity on a daily basis. In long term experiments on food utilization and growth rates using artificial diet, no beneficial effects of rutin could be demonstrated. Ingested rutin was absorbed and some of it deposited in the cuticle. Most was excreted as the aglycone quercetin. The possible functional significance of the phagostimulatory effect is discussed.Dedicated to G.B. Marini-Bettolo     

The relative insensitivity of Manduca sexta larvae to non-host plant secondary compounds

We tested the deterrent effects of non-host plant secondary compounds on Manduca sexta Johan. larvae, in choice tests. Treatment chemicals were applied, at approximately natural concentrations to either glass fiber or tobacco leaf disks. Of 30 compounds tested on glass fiber disks, with artificial diet reared larvae, 6 were deterrent. When the 6 deterrent chemicals were tested on tobacco leaf disks only one was still deterrent. No deterrent effects were observed when tobacco reared larvae were tested with chemicals applied to tobacco leaf disks. The results suggest that maintenance of restricted host range in M. sexta larvae relies not on avoidance of non-hosts due to deterrence but on attraction and stimulation to feed on hosts.

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Résumé Nous avons examiné au moyen des expériences de choix, les effects dissuasifs des composés secondaires de plantes non-hôtes des chenilles de M. sexta Johan. Des disques de fibres de verre ou des rondelles de feuilles de tabac ont été traités chimiquement, aux concentrations à peu près naturelles. Sur les 30 substances examinées sur les disques de fibres de verre, 6 ont été répulsives pour des chenilles élvées sur aliment artificiel. Quand l'examen est effectué avec des rondelles de feuilles de tabac, seul un produit est encore répulsif. Aucun effet répulsif n'est observé quand des chenilles élevées sur tabac sont mises en présence de substances associées à des rondelles de feuille de tabac. Ces résultats suggèrent que le maintien d'un spectre d'hôtes limité chez les chenilles de M. sexta ne repose pas sur l'évitement des plantes non-hôtes par suite de leurs substantes dissuasives, mais sur l'attraction et la stimulation provoquées par les hôtes.

     

Evaluation of acute and chronic toxicity of selected compounds in higher plants using cell culture

Summary The feasibility of using plant cell culture to measure toxicity was determined by investigating the toxicological effects of three chemical compounds, allyl alcohol, propargylglycine, and cadmium chloride, on cell cultures ofCatharanthus roseus G. Don (Madagascar periwinkle). Suspension cultures ofC. roseus were maintained in modified B5 medium and transferred every 5 d. Five-day-old cell cultures were exposed to various concentrations (10,3,1,0.3,0.1,0.03,0.01,0.003,0.001,0.0003,0.0001, 0.00003, and 0.0 mM) of the toxicants in both acute and chronic toxicity tests. In the acute test, cells were exposed to the toxicant for 24 h, washed three times with sterile medium, and plated in petri plates with an equal volume of 1.4% agar medium. Cells in the chronic test were plated with an equal volume of 1.4% agar medium containing various concentrations of the toxicant. Cells were incubated 28 d at 30°C in the dark. The colonies were counted and the results plotted as percent survival versus toxicant concentration. The results indicate, at the concentrations tested, thatC. roseus assay may be feasible in that it fulfills the criteria for a practical assay (e.g., rapid, simple, quantifiable, and reproducible). This work was submitted to the faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Environmental Science, Institute of Environmental Sciences.     

Relationship between deterrence and toxicity of plant secondary compounds for the alfalfa weevil Hypera brunneipennis

A variety of plant secondary compounds, several of which are quite widespread in nature were tested for their deterrence to the specialist coleopteran Hypera brunneipennis (Boheman) in short-term behavioral assays. The compounds were nicotine, quinine, sparteine, hordenine, linamarin, amygdalin, sinigrin, morin, juglone, chlorogenic acid, digitonin, mimosine, diosgenin, rutin and ursolic acid. Nine of these were then tested for their post-ingestional effects over one to two weeks of adult life, using fecundity as a measure of the effects. In only one case was there any indication of a detrimental effect or any trend suggesting one. The evolutionary implications of these findings are discussed.     

Food acceptance by a monophagous and an oligophagous insect in relation to seasonal changes in host plant suitability

Food acceptance by larvae of two lepidopteran species feeding on Rosaceae, viz. Yponomeuta evonymellus (monophagous) and Y. padellus (oligophagous), was compared. The influence of seasonal changes in plants as food for both insects was examined, in particular, the effects of nitrogen and sorbitol in leaves. In the laboratory, Y. evonymellus accepts Crataegus monogyna, a host plant of Y. padellus, and Y. padellus accepts Prunus padus, the host plant of Y. evonymellus. P. padus is the most suitable food plant for Y. evonymellus. No difference in food-quality for Y. padellus was found between C. monogyna and P. padus. The performance of both species on P. padus is less influenced by seasonal changes than on Crataegus. The suitability of Crataegus decreases during the season. This is probably caused by the decrease of its nitrogen content, and not by the decrease of sorbitol in the plant. The monophagous, Y. evonymellus, is more sensitive to seasonal changes in its food when fed with a non host plant than the oligophagous Y. padellus. In oviposition experiments both species have a preference for their normal host-plants.     

Food mixing by generalist grasshoppers: Plant secondary compounds structure the pattern of feeding

Plant secondary compounds can prevent feeding by phytophagous insects or, if ingested, can be poisonous to them. Less attention has been paid to the additional effects they have on feeding behavior when they are only weakly deterrent or not deterrent at all. Experiments were carried out on the generalist grass-hopperSchistocerca americana. Individuals were presented either with two cakes of high-quality artificial food with a single deterrent compound added or with two cakes, each with a different added deterrent compound. The deterrents consisted of single plant secondary compounds that were either marginally or strongly deterrent. There were profound differences in feeding behavior between those individuals given identical and those given different cake types, including longer feeding bouts on single cakes when a choice of different cake types was available. The behavioral effects demonstrate that the presence of secondary compounds in one food can influence the patterns of feeding on other available foods and suggest that such chemicals could impact foraging activities in a complex manner.     

南美斑潜蝇寄主选择性与植物次生化合物及叶毛的关系

采用非自由选择法,以刺伤孔（包括取食孔和产卵孔）数为指标,在室内测定了南美斑潜蝇对6科、16种、22个蔬菜品种的寄主选择性。结果表明,南美斑潜蝇对供试蔬菜品种的选择性存在显著差异,选择性大小依次为豆科>菊科>葫芦科>茄科>伞形科>十字花科; 菜豆和茼蒿是南美斑潜蝇最喜欢产卵和取食的寄主,而甘蓝和番茄是其选择性最差的寄主。化学分析和解剖镜观察表明,叶片单宁酸、黄酮含量及叶毛密度在供试品种间存在显著差异： 苦瓜的单宁酸含量最高,其次为豌豆、番茄和青椒,含量最低的为金丝搅瓜、茼蒿、菜豆和美国西芹; 黄酮含量最高的为甘蓝和番茄,其次为苦瓜,最低为茼蒿、菜豆和美国西芹; 南瓜和小白菜的叶毛密度最高,其次为西葫芦、金丝搅瓜和甘蓝,最低为青椒、莴苣和苦瓜。相关分析表明,南美斑潜蝇寄主选择性与叶片单宁酸和黄酮含量呈显著的负相关关系,相关系数分别为-0.4425（ P=0.0392）和 -0.5728（ P=0.0053）,而与叶毛密度相关关系不显著（ R＝-0.1807,P=0.4211）。说明黄酮和单宁酸对南美斑潜蝇产卵和取食具有明显的抑制作用。     

Larval growth inhibition of the spiny bollworm,Earias insulana, by some steroidal secondary plant compounds

The effect of 23 secondary plant compounds including a series of steroidal aglycones and glycosides, on larval growth and pupation ofEarias insulana (Boisd.) was investigated. Larvae did not develop when fed on artificial diets containing 0.2% solamargine, solasonine, tomatine, digitonin, saponin, nomilin, lawsone or coumarin. Some growth-retarding activity was found with diets containing 0.2% ajmalicine, capsaicin, quercetin, glycyrrhizin and glycerrhetic acid. The eight compounds highly active at 0.2% were also bioassayed at 0.1, 0.05 and 0.01%: larvae did not pupate when fed on diets containing 0.1% solasonine, tomatine or nomilin, whereas solamargine and coumarin were active even at 0.05%. Preliminary structure-activity relationships were evaluated for the active steroidal glycosides.

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Larvale Wachstumshemmung des ägyptischen Baumwollkapselwurms, Earias insulana,durch einige steroide sekundäre Planzeninhaltstoffe

Zusammenfassung Es wurde die Wirkung von 23, einem künstlichen Nährmedium einverleibten, sekundären Pflanzeninhaltstoffen, unter ihnen eine Reihe steroider Aglucone und Glycoside, auf das Larvenwachstum und die Verpuppung vonEarias insulana untersucht. Larven entwickelten sich nicht in mit 0.2% Solamargin, Solasonin, Tomatin, Digitonin, Saponin, Nomilin, Lawsone und Coumarin versetzten Nährmedien. Auf Medien, die 0.2% Ajmalicin, Capsaicin, Quercetin, Glycyrrhizin and Glycyrrhetinsäure enhielten, war das Larvenwachstum verzögert. Die 8 bei 0.2% hoch wirksamen·Substanzen wurden auch bei 0.1, 0.05 und 0.01% getestet. Die Larven verpupten sich nicht auf 0.1% Solasonin, Tomatin und Nomilin und auf 0.05% Solamargin und Coumarin. Vorläufige Struktur-Wirksamkeit-Beziehungen wurden für die aktiven steroiden Glycoside vorgeschlagen.

     

Habituation in Frankliniella occidentalis to deterrent plant compounds and their blends

Feeding and oviposition deterrence of three secondary plant compounds and their 1:1 blends to adult female Frankliniella occidentalis Pergande (Thysanoptera: Thripidae) and the potential for habituation of the thrips to the pure compounds and the 1:1 blends at various concentrations were investigated. In choice assays, we tested dose‐dependent feeding and oviposition deterrence of the two fatty acid derivatives methyl jasmonate and cis‐jasmone, the phenylpropanoid allylanisole, and their blends when directly applied to bean leaf discs. The concentration required to reduce the feeding damage by 50% relative to the control treatment (FDC₅₀) was lowest for cis‐jasmone and highest for allylanisole. The feeding deterrent effect of both jasmonates was increased when blended with allylanisole. Feeding deterrence and oviposition deterrence were strongly correlated. In no‐choice assays conducted over four consecutive days, we discovered that dilutions at low concentrations (FDC₁₅) applied to bean leaves resulted in habituation to the deterrents, whereas no habituation occurred at higher concentrations (FDC₅₀). We observed a tendency that the 1:1 blends reduce the probability that thrips habituate to the deterrent compounds. Our results may be useful in the development of integrated crop protection strategies with the implementation of allelochemicals as pest behaviour‐modifying agents.     

Interactions of plant stress and herbivory: intraspecific variation in the susceptibility of a palatable versus an unpalatable seaweed to sea urchin grazing

Summary Interactions among environmental stresses, plant defensive characteristics, and plant nutrient status may significantly affect an alga's susceptibility to herbivores. Following desiccation, the palatable seaweed Gracilaria tikvahiae was less susceptible to grazing by the sea urchin Arbacia punctulata while the unpalatable alga Padina gymnospora became more susceptible. Increased grazing on desiccated Padina appeared to result from a loss of chemical defenses following desiccation. Palatable plants treated with organic extracts from desiccated Padina plants were consumed at more than twice the rate of plants treated with extracts from undesiccated plants. Increased susceptibility of Padina did not correlate with changes in protein content of the alga; reduced grazing on desiccated Gracilaria was associated with a decrease in protein content. When Padina was grazed by Arbacia or mechanically damaged to mimic urchin grazing, its susceptibility to Arbacia decreased within 1 to 5 days. These results demonstrate that history of physical or biological stress may affect a plant's susceptibility to herbivory. We hypothesize that urchins cue primarily on attractiveness features (e.g. nutrient content) of highpreference algae and deterrent features (e.g. chemical defenses) of low-preference algae. Stresses may therefore increase, decrease, or not affect a plant's susceptibility to herbivory depending upon the primary feeding cues used by the herbivore, the defensive mechanisms used by the plant, and the way these are altered by various environmental stresses.     

Effect of plant secondary compounds on in vitro methane,ammonia production and ruminal protozoa population

Aims

The objective of this study was to evaluate the potential of secondary plant metabolites from 38 sources to serve as antimethanogenic additives in ruminant diets. The effect of leaf tannins from these different plant sources on rumen fermentation, protozoal populations and methanogenesis was also studied.

Methods and Results

Samples (200 mg dry matter, DM) were incubated without and with polyethylene glycol (PEG)‐6000 (400 mg DM) as a tannin binder during 24‐h incubation in the in vitro Hohenheim gas system. In the leaf samples, total phenol (g kg^?1 DM) was maximum in Pimenta officinalis (312) followed by Oenothera lamarckiana (185) and Lawsonia inermis (105). Of the 38 samples, condensed tannins exceeded 4·0 g kg^?1 in only Alpinia galanga (7·50), Cinnamomum verum (4·58), Pelargonium graveolens (18·7) and Pimenta officinalis (23·2) and were not detected in seven samples. When the bioactivity of the leaf samples was assessed using the tannin bioassay, the percentage increase in the amount of gas produced during incubation of samples with the tannin‐binding agent PEG‐6000 over the amount produced during incubation without the tannin binder ranged from nil (zero) to 367%, with the highest being recorded with A. galanga leaves. The ratio of methane reduction per ml of total gas reduction was maximum with Rauvolfia serpentina (131·8) leaves, followed by Indigofera tinctoria (16·8) and Withania somnifera (10·2) leaves. Total and differential protozoal counts increased with added PEG in twenty‐two samples, maximum being in Pimenta officinalis. Increased accumulation of total volatile fatty acids during incubation with added PEG‐6000 was recorded, and the values ranged from zero to 61%. However, the increase was significant in only 11 of the 38 tannin sources tested indicating noninterference of tannin on in vitro fermentation of carbohydrates by the majority of samples tested. Conversely, in 26 of 38 plant sources, the leaf tannins reduced N‐digestibility as evidenced by increased accumulation of NH₃‐N with added PEG.

Conclusions

Our study unequivocally demonstrated that plants containing secondary metabolites such as Rauvolfia serpentine, Indigofera tinctoria and Withania somnifera have great potential to suppress methanogenesis with minimal adverse effect of feedstuff fermentation.

Significance and Impact of the Study

It was established that methanogenesis was not essentially related to the density of protozoa population in vitro. The tannins contained in these plants could be of interest in the development of new additives in ruminant nutrition.     

The effect of variation in host plant on the growth of an oligophagous insect, Malacosoma americanum and its polyphagous relative, Malacosoma disstria

Larvae of Malacosoma americanum (F.)(Lepidoptera: Lasiocampidae) an oligophagous species that feeds primarily on Prunus and other rosaceous trees, were compared to larvae of the more highly polyphagous congener M. disstria Hb., with respect to their sensitivity to variation in the foliage of a common host plant, Prunus serotina Ehrh. Pupal weight and time to pupation were measured on larvae reared on foliage from open-grown and from shaded saplings. The difference in foliage had a pronounced effect, but no difference was evident between the species in their response to the variation in foliage. This comparison implies that sensitivity to intraspecific variation in host quality does not differ between host-specific and generalized species. However, results from other species suggest that some species of insects do differ in this respect.

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Zusammenfassung Raupen von Malacosoma americanum (F.), einer oligophagen Art, die sich vor allem auf Prunus und andern baumartigen Rosaceen entwickelt, wurden verglichen mit Raupen der polyphageren Verwandten M. disstria Hb. und zwar im Hinblick auf deren Empfindlichkeit auf Unterschiede im Blatt ihrer gemeinsamen Wirtspflanze, Prunus serotina Ehrh. Das Puppengewicht und die Entwicklungszeit bis zur Verpuppung wurden gemessen bei Raupen, welche auf Blättern von freiwachsenden und von beschatteten Jungpflanzen gezüchtet worden waren. Die Blattunterschiede hatten eine ausgesprochene Wirkung, aber es gab keine Unterschiede in der Reaktion der beiden Arten. Dieser Vergleich lässt vermuten, dass die Empfindlichkeit auf intraspezifische Unterschiede der Wirtspflanzenqualität bei wirtsspezifischen und polyphagen Arten gleich ist. Indessen dürften sich laut anderen Resultaten einige Insektenarten anders verhalten.

     

Influence of an endophytic fungus on host plant selection by a polyphagous moth via volatile spectrum changes

Recently the role of micro-organisms as mediators of plant-herbivore interactions has been increasingly acknowledged in ecological research. We investigated the interaction between an unspecialized root fungal endophyte (Acremonium strictum) and the polyphagous moth Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), in greenhouse and laboratory bioassays. Specifically we examined in tomato (Lycopersicon esculentum Mill.) the systemic effects of the endophytic fungus on the host selection behaviour of female moths for oviposition and the volatile profiles of host plants to understand the mechanisms acting in this multi-trophic model system. Both laboratory and field strains of H. armigera moths oviposited more on leaves of A. strictum inoculated plants as compared to endophyte-free plants, both in free flight cages and in tethered moth laboratory experiments; the moth’s preferences were significant between 10 and 18 days after inoculation. The analysis of volatile profiles showed strong quantitative differences between treatments. Endophyte inoculated plants emitted diverse terpenes and sesquiterpenes at significantly lower amounts as compared to endophyte free-plants, except for α-terpinene, which did not differ between the treatments, and trans-β-caryophyllene, which was emitted in significantly higher amounts on inoculated plants. β-Thujene and α-phellandrene accounted for 73.3 and 12.0% of total amounts of volatiles emitted from endophyte-free and inoculated plants, respectively. Our findings demonstrate that A. strictum is able to systemically influence the host selection of H. armigera moths for oviposition; conceivably mediated by the induced changes in volatile emissions (and probably additional biochemical parameters of the host plants, which have not been analysed so far). We argue for a more detailed assessment of micro-organisms invisibly colonizing plants, when studying plant-herbivore or multitrophic interactions.     

Indirect interaction between an unspecialized endophytic fungus and a polyphagous moth

Despite their ubiquitous occurrence in natural and managed ecosystems, the role of unspecialized fungal endophytes in mediating insect-plant interactions have been largely ignored compared to clavicipitaceous fungal endophytes of grasses. We conducted laboratory and glasshouse experiments to test the hypothesis that an unspecialized fungal endophyte (Acremonium strictum) restricted to the root system of the host plant is able to mediate the interaction between a polyphagous herbivorous insect (Helicoverpa armigera) and its host plant (tomatoes, Lycopersicum esculentum). Our studies indicated a significant variation in the ability of inoculated and control plants to support the life history stages of the insect. Larvae reared on inoculated plants suffer significant reduction in growth rate, prolonged development times, suppressed moulting, and produced smaller pupae that were more likely to die, and emerged adults were less fecund compared to larvae reared on control plants. In glasshouse bioassays were larvae were allowed to graze freely, on inoculated plants only 20% survived to the final stadium compared to 54.5% on control plants. No significant differences in the amount of foliage consumed were found between inoculated and control plants, suggesting that A. strictum does not act as a feeding deterrent for H. armigera larvae. Larvae fed inoculated plants showed a significant increase in the relative consumption rate, although the approximate digestibility of ingested food, and efficiency with which both ingested food and digested food was converted to insect biomass decreased during the same period. In addition to reduced efficiency of food utilization, surviving larvae on inoculated plants displayed a significant reduction in relative growth rate. We hypothesized that alterations in phytosterol composition mediated by infection of the fungal endophyte, may explain the reduced larval performance on inoculated plants.Obwohl unspezialisierte endophytische Pilze in allen Ökosystemen weit verbreitet und häufig sind, blieb ihre Rolle bei Interaktionen zwischen Pflanzen und herbivoren Insekten bisher weitgehend unberücksichtigt. In Labor- und Gewächshausexperimenten haben wir die Hypothese untersucht, dass ein unspezialisierter Wurzelendophyt (Acremonium strictum) die Wechselwirkungen zwischen einem polyphagen Insekt (Helicoverpa armigera) und Tomatenpflanzen (Lycopersicum esculentum) beeinflussen kann. Die Entwicklung der Insekten weist deutliche Unterschiede zwischen inokulierten und Kontrollpflanzen auf. Larven, die auf inokulierten Pflanzen gehalten wurden, zeigten gegenüber solchen von Kontrollpflanzen jeweils signifikant verringerte Wachstumsraten, verlängerte Entwicklungszeiten, verzögerte Häutungen, und bildeten kleinere Puppen aus, welche eine höhere Mortalität aufwiesen. Die geschlüpften Weibchen hatten zudem eine verringerte Fekundität. In Frassversuchen im Gewächshaus überlebten an inokulierten Pflanzen nur 20% der Larven während an Kontrollpflanzen 54,5% überlebten. Da die gefressene Biomasse in beiden Varianten gleich war, ist eine deterrente Wirkung des Pilzes auszuschließen. Vielmehr deuten signifikante Veränderungen der Fraß- und Verwertungsindices sowie die niedrigeren relativen Wachstumsraten darauf hin, das sich die Nahrungsqualität auf inokulierten Pflanzen verschlechtert hatte. Wir diskutieren die Hypothese, dass der Phytosterolgehalt, welcher durch eine Inokulation der Pflanzen verändert wird, zu den verschlechterten Entwicklungsbedingungen der Larven geführt hat.     

Susceptibility and intrinsic tolerance of Pseudomonas aeruginosa to selected plant volatile compounds

AIMS: To examine the causes for variations in sensitivity and intrinsic tolerance of Pseudomonas aeruginosa to plant volatile compounds. METHODS AND RESULTS: Minimum inhibitory concentrations were determined for a selection of volatile phytochemicals against P. aeruginosa using a microdilution assay. Effects on growth were also assessed in 100-ml broth cultures. The two strains of P. aeruginosa included in the study exhibited intrinsic tolerance to all compounds, with the exception of carvacrol and trans-cinnamaldehyde. The protonophore carbonyl cyanide m-chlorophenylhydrazone increased P. aeruginosa sensitivity to all compounds except trans-cinnamaldehyde, implicating an ATP-dependent efflux mechanism in the observed tolerance. Outer membrane integrity following treatment with test compounds was assessed by measuring sensitization to detergents. Only carvacrol caused damage to the outer membrane of P. aeruginosa. CONCLUSIONS: The intrinsic tolerance of P. aeruginosa strains to plant volatile compounds is associated with an active efflux mechanism and the barrier function of the outer membrane. SIGNIFICANCE AND IMPACT OF THE STUDY: These findings offer an explanation for the intrinsic tolerance to plant volatile compounds exhibited by P. aeruginosa. The study also confirms that the outer membrane-permeabilizing action of carvacrol, previously reported for the gram-negative bacteria Escherichia coli and Salmonella, extends to monoterpene-tolerant strains of P. aeruginosa.     

The toxicity of phenolic compounds to soil algal population and toChlorella vulgaris andNostoc linckia

Summary p-Nitrophenol (PNP),m-nitrophenol (MNP), 2,4-dinitrophenol (DNP) and catechol were tested for their effects on algal population in a soil and on pure cultures of two algae isolated from soil. Both PNP and MNP, even at 0.5 kg ha⁻¹ level were toxic to the soil algae; high doses effected increase in toxicity. Inhibition of algae was relatively more with PNP compared to the other two nitrophenols. Catechol treatment up to 1.0 kg ha⁻¹ led to a significant initial enhancement of algae with a subsequent far less toxic effect. The toxicity of the phenolic compounds towardChlorella vulgaris, a green alga andNostoc linckia, a blue-green alga, decreased in the order: MNP≧PNP>DNP>Catechol. However, algicidal or algistatic effect of the test chemicals was fairly more againstC. vulgaris, suggesting that the eukaryotic alga is highly sensitive to such soil pollutants compared to the prokaryotic alga.     

How do herbivorous insects cope with noxious secondary plant compounds in their diet?

Herbivorous insects use a variety of physiological mechanisms to cope with noxious (i.e., unpalatable and/or toxic) compounds in their food plants. Here, I review what is known about this coping process, focusing on one species of caterpillar, the tobacco hornworm (Manduca sexta). Herbivorous insects possess both preingestive (i.e., chemosensory) and postingestive response mechanisms for detecting plant secondary compounds. Stimulation of either class of detection mechanism inhibits feeding rapidly by reducing biting rate and/or bite size. This aversive response is highly adaptive during encounters with secondary plant compounds that are toxic. The insect's dilemma is that many harmless or mildly toxic compounds also activate the aversive response. To overcome this dilemma, herbivorous insects employ at least three mechanisms for selectively deactivating their aversive response to relatively harmless secondary plant compounds: (1) the presence of carbohydrates can mask the unpalatable taste of some secondary plant compounds; (2) prolonged dietary exposure to some unpalatable secondary plant compounds can initiate long-term adaptation mechanisms in the peripheral and central gustatory system; and (3) dietary exposure to toxic compounds can induce production of P450 detoxication enzymes. Thus, herbivorous insects utilize an integrated suite of physiological mechanisms to detect potentially toxic compounds in foods, and then selectively adapt to those that do not pose a serious threat to their growth and survivorship.     

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.