Differential predation by Coleomegilla maculata on Colorado potato beetle strains that vary in growth on tomato

This study tests the hypothesis that the generalist predator Coleomegilla maculata DeGeer causes differential mortality of Colorado potato beetle, Leptinotarsa decemlineata (Say), larvae differing in their degree of genetic adaptation to tomato (Lycopersicon esculentum Mill.) as a host plant. Results of a series of laboratory experiments demonstrate that adult C. maculata can cause higher mortality to nonadapted than adapted Colorado potato beetle larvae. The extent of differential mortality caused by C. maculata depended on age of potato beetle larvae; presence of potato beetle eggs; whether or not the predator had a choice among prey items; and, in choice situations, the ratio of adapted to nonadapted potato beetle larvae. Although adult C. maculata have the potential to prey differentially on tomato-adapted and nonadapted Colorado potato beetle larvae in mixed populations, the magnitude of differential predation in a natural setting could be highly variable.     

Comparative headspace analysis of cabbage plants damaged by two species ofPieris caterpillars: consequences for in-flight host location byCotesia parasitoids

Headspace composition, collected from intact cabbage plants and cabbage plants infested with eitherPieris brassicae L. orP. rapae L. (Lepidoptera: Pieridae) first instar larvae, was determined by GC-MS. Twenty-one volatiles were identified in the headspace of intact plants. Twenty-two volatiles were identified in the headspace of plants infested byP. brassicae larvae, 2 of which, Z-3-hexenyl butyrate and Z-3-hexenyl isovalerate, were not detected in the headspace of either intact orP. rapae damaged plants. In the headspace of the latter, 21 compounds were identified, all of which which were also produced by intact plants. No significant quantitative differences were found between headspace composition of the plants damaged by one or the other caterpillar species. Major differences between intact and caterpillar-damaged plants in contribution to the headspace profile were revealed for hexyl acetate, Z-3-hexenyl acetate, myrcene, sabinene and 1,8-cineole. The larval endoparasitoidCotesia glomerata L. was attracted by the volatiles emanating fromB. oleracea damaged byP. brassicae first instar larvae.C. rubecula L., a specialized larval endoparasitoids ofP. rapae, was attracted by the volatiles released from theB. oleracea-P. rapae plant-host complex. This shows that cabbage plants kept under the conditions of headspace collection produce attractive volatiles for both parasitoids.     

Volatiles released from bean plants in response to agromyzid flies

Liriomyza sativae Blanchard and Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) are two invasive flies in China that have caused economical damage on vegetables and ornamental plants. In this article, we report the profiles of emitted volatiles from healthy, mechanically damaged, and leafminer-damaged bean, Phaseolus vulgaris L., plants. Among 25 emitted volatiles identified, (E)-2-hexen-1-al, (3E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, (syn)- and (anti)-2-methylpropanal oxime, (syn)-2-methylbutanal oxime, linalool, and (E,E)-α-farnesene were consistently released from damaged bean plants. Combined amounts of these nine compounds made up more than 70% of the total volatiles emitted from each treatment. No qualitative differences in volatile emission were found between bean plants damaged by the two fly species; however, amounts of several major compounds induced by L. huidobrensis damage were significantly higher than those from plants damaged by L. sativae. The mechanically damaged plants released a higher proportion of green leaf volatiles than plants in the other treatments, whereas leafminer-damaged plants produced more terpenoids and oximes. Furthermore, the volatile profiles emitted from plants, damaged by adult leafminers, by second instar larvae, and even the plants with empty mines left by leafminer larvae (the pupal stage) were significantly different. The identification of volatile oximes released from damaged plants was confirmed and is discussed in a behavioral and biological control context.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Oriented responses of the triatomine bugs Rhodnius prolixus and Triatoma infestans to vertebrate odours on a servosphere

Oriented responses of both R. prolixus and T. infestans adults were recorded on a servosphere to mouse-odour, one of its components (CO₂), and to rabbit urine-odour. The volatiles were delivered in an air-stream under controlled conditions which excluded other sensory modalities. In stimulus-free air the triatomines walked preferentially downwind in straight bouts interrupted by stops or periods at relatively low speeds, all of variable duration. In odour-laden air, bugs maintained their typical walking habit but switched from negative to positive anemotaxis. The characteristic response to odour onset was to stop, sample the air with the antennae, turn upwind in situ, and then walk off in the direction of the source for at least a few seconds, i.e., odour mediated anemotaxis. Mouse-odour caused T. infestans to increase its speed to 5.3 cms^-1. Both species continued with the upwind response for some time after odour delivery ceased, but the crosswind component of the tracks was more prominent during this period — an effort, we presume, by the bugs to re-contact an odour plume. This investigation provides unequivocal evidence for host finding in triatomines by olfactory cues alone.     

Can herbivore‐induced plant volatiles inform predatory insect about the most suitable stage of its prey?

The leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) is a specialist herbivore, all of whose mobile stages feed on the leaves of salicaceous plants. Both the larval and adult stages of the ladybird Aiolocaria hexaspilota (Coleoptera: Coccinellidae) are dominant natural enemies of the larvae of the leaf beetle. To clarify the role of plant volatiles in prey‐finding behaviour of A. hexaspilota, the olfactory responses of the ladybird in a Y‐tube olfactometer are studied. The ladybird adults show no preference for willow plants Salix eriocarpa that are infested by leaf beetle adults (nonprey) over that for intact plants but move more to the willow plants infested by leaf beetle larvae (prey) than to intact plants. Moreover, ladybird larvae show no preference for willow plants infested by leaf beetle larvae or adults over intact plants. Using gas chromatography‐mass spectrometry, six volatile compounds are released in larger amounts in the headspace of willow plants infested by leaf beetle larvae than in the headspace of willow plants infested by leaf beetle adults. In addition, the total amount of volatiles emitted from willow plants that are either intact or infested by leaf beetle adults is much smaller than that from willow plants infested by leaf beetle larvae. These results indicate that volatiles from S. eriocarpa infested by P. versicolora inform A. hexaspilota adults about the presence of the most suitable stage of their prey, whereas A. hexaspilota larvae do not use such information.     

Variation in damage to cotton affecting larval feeding preference of Spodoptera littoralis

Feeding experiments with larvae of Spodoptera littoralis were performed with leaves from cotton plants subjected to damage and from undamaged plants. In the experiments, four different time intervals (1, 3, 7, and 14 days) after damage induction and two different levels (high and low) of herbivore damage were tested. Seven days after damage induction larvae fed less on the young top leaves from damaged plants for both levels of damage. At the high damage level, the larvae fed less on leaves from the damaged plants after just three days, and this effect still remained 14 days after damage infliction. When mature leaves from the middle of the plant were compared, no difference between treatments was observed.Two plant sizes were tested, small plants with 4–5 true leaves and large plants with 8–10 true leaves. In small plants the induced changes affecting larval feeding were found mainly in the youngest leaf at the top of the plant, while in large plants the induced effects were found in both the youngest and the second youngest leaves.In plants subjected to artificial damage, larvae fed less on top leaves of the damaged plants when compared to leaves from undamaged plants. When leaves from plants that had been artificially damaged were directly compared with leaves from plants damaged by herbivores, larvae fed more on the youngest leaves from artificially damaged plants when the plants were large. In small plants no significant difference was found when comparing artificial and herbivore damage.     

Response to host plant odors and aggregation pheromone by larvae of the Colorado potato beetle on a servosphere

Using a servosphere (locomotion compensator), locomotory behavior of Colorado potato beetle Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae) larvae was measured in detail in response to pulsed and non-pulsed odors of hostplant and conspecific pheromone. Second instars showed decreased straightness of movement, and all larvae showed decreased speed, in response to potato odor. Change in straightness by 2nd instars was also significantly affected by the interaction of pheromone and pulsing treatments. Fourth instars showed increased positive anemotaxis in response to the combined hostplant and pheromone odors. To our knowledge, this is the first demonstration of larval behavioral responses to adult pheromone in a holometabolous insect.     

Characterization of induced resistance in tomato plants

We have characterized, using several types of bioassays, the resistance induced in young tomato plants by feeding of the corn earworm, Helicoverpa zea. Beet armyworm larvae, Spodoptera exigua, and leafminers, Liriomyza trifolii, were used to assay the induced resistance. In whole-plant experiments, damage localized to a single leaflet of fourleaf tomato plants induced a systemic increase in resistance such that beet armyworm larvae confined to previously damaged (induced) plants grew at a rate about half that of larvae raised on control plants and consumed less leaf tissue from induced plants than from control plants. In experiments using excised leaves, beet armyworm larvae suffered increased mortality when reared on leaves from induced plants. The strength of this induced resistance varied spatially relative to the damaged position; moreover, the spatial distribution of induced resistance changed over a three-week period following damage. Other experiments demonstrated that the mechanisms of induced resistance in tomato foliage involves both a decrease in larval preference for and a decrease in the nutritional value of induced foliage. Induction also retarded the oviposition and/or early development of leafminers. Thus, induced resistance has relatively severe effects on the biology of subsequent herbivores. These data should allow us to begin to elucidate cause-effect relationships between induced resistance and induced chemistry in tomato plants.     

The influence of host plant species on parasitism of pollen beetles (Meligethes spp.) by Phradis morionellus

We investigated the response of the specialist insect predator Oligota kashmirica benefica (Coleoptera: Staphylinidae) to volatiles from lima bean leaves infested with the spider mite Tetranychus urticae (Acari: Tetranychidae), both in a Y-tube olfactometer and in a field in Kyoto, Japan. Adult male and female predators were significantly more attracted to T. urticae-infested leaves than to clean air. Adult male and female predators were not more attracted to uninfested leaves, artificially damaged leaves, or the spider mites and their visible products when compared to clean air. In a field trap experiment, 12 adult predators were caught in three traps containing T. urticae-infested lima bean plants over 13 days, whereas no adult predators were trapped in three traps containing uninfested lima bean plants during the same period. These results showed that O. kashmirica benefica adults responded to herbivore-induced plant volatiles from T. urticae-infested lima bean leaves under both laboratory and field conditions.     

Parasitoid-plant mutualism: parasitoid attack of herbivore increases plant reproduction

We tested whether a plant's life time seed production is increased by parasitization of herbivores in a tritrophic system, Arabidopsis thaliana (Brassicaceae) plants, Pieris rapae (Lepidoptera: Pieridae) caterpillars and the solitary endoparasitoid Cotesia rubecula (Hymenoptera: Braconidae). We established seed production for intact A. thaliana plants, plants that were mechanically damaged, plants fed upon by parasitized caterpillars and plants fed upon by unparasitized caterpillars. In the first experiment, with ecotype Landsberg (erecta mutant), herbivory by unparasitized P. rapae caterpillars resulted in a strongly reduced seed production compared to undamaged plants. In contrast, damage by P. rapae caterpillars that had been parasitized by C. rubecula did not result in a significant reduction in seed production. For the second experiment with the ecotype Columbia, the results were identical. Plants damaged by unparasitized caterpillars only produced seeds on regrown shoots. Seed production of plants that had been mechanically damaged was statistically similar to that of undamaged plants. Production of the first ripe siliques by plants fed upon by unparasitized caterpillars was delayed by 18–22 days for Landsberg and 9–10 days for Columbia. We conclude that parasitization of P. rapae by C. rubecula potentially confers a considerable fitness benefit for A. thaliana plants when compared to plants exposed to feeding damage by unparasitized P. rapae larvae. Plants that attract parasitoids and parasitoids that respond to herbivore-induced plant volatiles will both experience selective advantage, justifying the use of the term mutualism for this parasitoid-plant interaction. This type of mutualism is undoubtedly very common in nature.     

Response of Neoseiulus fallacis Garmen and Galendromus occidentalis Nesbitt (Acari: Phytoseiidae) to Tetranychus urticae Koch (Acari: Tetranychidae)-Damaged Hop Humulus lupulus (L.) (Urticales: Cannabaceae)

Abstract 1 The response of Neoseiulus fallacis Garmen and Galendromus occidentalis (Acari: Phytoseiidae) to Tetranychus urticae Koch (Acari: Tetranychidae)‐damaged and undamaged hop, Humulus lupulus (L.), plants was tested using a Y‐tube olfactometer. 2 Neoseiulus fallacis but not G. occidentalis was attracted to volatiles from T. urticae‐damaged hop plants when paired with undamaged plants. 3 The response of N. fallacis to these volatiles was stronger for plants severed at the soil surface than for intact plants. 4 There was no difference in the response of N. fallacis to severed or intact hop plants that had no spider mite damage, indicating that artificial wounding by severing alone does not elicit the production of attractive volatiles detectable to N. fallacis. These results are consistent with the existence of cross‐talk between signalling pathways initiated by feeding damage and artificial wounding that result in elevated levels of predator‐attracting volatiles.     

Feeding damage by larvae of the mustard leaf beetle deters conspecific females from oviposition and feeding

Herbivorous insects may be informed about the presence of competitors on the same host plant by a variety of cues. These cues can derive from either the competitor itself or the damaged plant. In the mustard leaf beetle Phaedon cochleariae (Coleoptera, Chrysomelidae), adults are known to be deterred from feeding and oviposition by the exocrine glandular secretion of conspecific co-occurring larvae. We hypothesised that the exocrine larval secretion released by feeding larvae may adsorb to the surface of Chinese cabbage leaves, and thus, convey the information about their former or actual presence. Further experiments tested the influence of leaves damaged by conspecific larvae, mechanically damaged leaves, larval frass and regurgitant on the oviposition and feeding behaviour of P. cochleariae. Finally, the effect of previous conspecific herbivory on larval development and larval host selection was assessed. Our results show that (epi)chrysomelidial, the major component of the exocrine secretion from P. cochleariae larvae, was detectable by GC-MS in surface extracts from leaves upon which larvae had fed. However, leaves exposed to volatiles of the larval secretion were not avoided by female P. cochleariae for feeding or oviposition. Thus, we conclude that secretion volatiles did not adsorb in sufficient amounts on the leaf surface to display deterrent activity towards adults. By contrast, gravid females avoided to feed and lay their eggs on leaves damaged by second-instar larvae for three days when compared to undamaged leaves. Mechanical damage of leaves and treatment of artificially damaged leaves with larval frass or regurgitant did not affect oviposition and feeding of P. cochleariae. Since no adverse effects of previous herbivory on larval development were detected, we suggest that female P. cochleariae avoid Chinese cabbage leaves damaged by feeding larvae for other reasons than escape from competition or avoidance of direct negative effects that result from consuming induced plant material.     

Analyses of Two Parasitoids with Convergent Foraging Strategies

We compared the foraging strategies of two key braconid endoparasitoids of the tobacco budworm (Heliothis virescens Fab.), Cardiochiles nigriceps Vier. and Microplitis croceipes Cresson, that differ in host and habitat range but otherwise share comparable, overlapping niches. The most important host-location cues by far for both species were materials associated with damaged plants. Both species demonstrated a significant preference for volatiles released from plants damaged by H. virescens larvae over those released from undamaged tobacco and cotton plants. In choice experiments with damaged tobacco versus cotton, M. croceipes showed a significant preference for cotton plants. In contrast, C. nigriceps preferred damaged tobacco plants. Plant compounds provoked a strong response even when released from systemically induced plants (from which damaged leaves, host, and host by-products were removed). C. nigriceps appears to have a much keener ability to locate hosts over long distances than M. croceipes. This observation may be related to the highly specialized nature of this parasitoid. The possible adaptive significance of the foraging behaviors of these two parasitoids is discussed.     

Effect of needle damage on the release rate of Masson pine (<Emphasis Type="Italic">Pinus massoniana</Emphasis>) volatiles

The effect of needle damage on the release rate of Masson pine (Pinus massoniana Lamb.) volatiles was examined. Needles were continuously damaged by mechanical damage (MDP) or by feeding of pine caterpillar (Dendrolimus punctatus) larvae (LFP); undamaged pine was used as a control (UDP). Volatiles were collected before damage, and at 16, 24, 40, 48, 64, 72, 88 and 96 h post-damage, and analyzed. The analyses revealed that 19 compounds identified as constitutive volatiles from UDP were terpenes and green leaf odors. The release rate of volatiles from MDP or LFP was higher than that from UDP. At 96 h post-damage, emission from MDP or LFP returned to the same level as that of UDP. Some volatiles, including sabinene, ocimene, limonene-1,2-epoxide, linalool, linalool acetate, germacrene d-4-ol, farnesol, and (E)-4,8-dimethyl-1,3,7-nonatriene were induced by mechanical damage and/or larval attack. Furthermore, the release rate of linalool acetate, farnesol, or (E)-4,8-dimethyl-1,3,7-nonatriene from LFP was higher than that from MDP. Based on an exact estimation of the proportion of damaged pine needles, a significant linear correlation between the release rate of total volatiles identified and the proportion of damaged needles was found in the case of LFP but not MDP.     

Intraspecific Variation in Chemical Attraction of Rice to Insect Predators

The olfactory response of predators of the brown planthopper,Nilaparvata lugensStål, to different genotypes of rice (14 cultivars and breeding lines ofOryza sativaL. and 1 wild species,Oryza nivaraSharma et Shastry) was measured in an airflow olfactometer. Odor from rice plants attracted more females of the mirid predatorCyrtorhinus lividipennisReuter than plain air (control) on only 6 of the 15 rice genotypes. Orientation ofC. lividipennistoward volatiles of certain rice genotypes was apparent even when the plants were free of the brown planthopper. However, the predator distinguished between prey-infested and uninfested plants and preferred plants with eggs over plants with nymphs. The predator did not distinguish different stages of plant growth (vegetative, booting, or flowering). Plants artificially injured to simulate brown planthopper oviposition wounds were not as attractive to the predator as plants on which the planthopper had oviposited. The preassay preconditioning on the cultivar TN1 did not produce a predator bias for this genotype. This suggests that rearing effects or chemically mediated associative learning reported for some natural enemies did not influenceC. lividipennis'host response. Results with another predator, the coccinellidMicraspis hirashimaiSasaji, produced less consistent behavior. Planthopper-infested plants attracted more females ofM. hirashimaithan unifested plants in only 1 of the 12 rice genotypes evaluated. Implications for augmenting predators by rice cultivar selection and modification are discussed.     

Predator and parasitoid attacking ant-attended aphids: effects of predator presence and attending ant species on emerging parasitoid numbers

Interaction between a predator and a parasitoid attacking ant-attended aphids was examined in a system on photinia plants, consisting of the aphid Aphis spiraecola, the two ants Lasius japonicus and Pristomyrmex pungens, the predatory ladybird beetle Scymnus posticalis, and the parasitoid wasp Lysiphlebus japonicus. The ladybird larvae are densely covered with waxy secretion and are never attacked by attending ants. The parasitoid females are often attacked by ants, but successfully oviposit by avoiding ants. The two ants differ in aggressiveness towards aphid enemies. Impacts of the predator larvae and attending ant species on the number of parasitoid adults emerging from mummies per aphid colony were assessed by manipulating the presence of the predator in introduced aphid colonies attended by either ant. The experiment showed a significant negative impact of the predator on emerging parasitoid numbers. This is due to consumption of healthy aphids by the predator and its predation on parasitized aphids containing the parasitoid larvae (intraguild predation). Additionally, attending ant species significantly affected emerging parasitoid numbers, with more parasitoids in P. pungens-attended colonies. This results from the lower extent of interference with parasitoid oviposition by the less aggressive P. pungens. Furthermore, the predator reduced emerging parasitoid numbers more when P. pungens attended aphids. This may be ascribed to larger numbers of the predator and the resulting higher levels of predation on unparasitized and parasitized aphids in P. pungens-attended colonies. In conclusion, a negative effect of the predator on the parasitoid occurs in ant-attended aphid colonies, and the intensity of the interaction is affected by ant species.     

Short-term damage-induced increases in tobacco alkaloids protect plants

Summary Leaf damage significantly increases the alkaloid content in undamaged leaves on damaged field-grown wild tobacco plants. Although field-grown pot-bound plants fail to exhibit the same damage-induced increase in alkaloid content, the ability to respond to leaf damage is restored 6 days after removing plants from their pots. Freshly hatched Manduca sexta larvae reared individually in the laboratory on the high-alkaloid foliage of damaged plants released from their pots gain less weight and eat less (57.2% and 45.7% of controls, respectively) than larvae fed low-alkaloid foliage from undamaged released plants. Moreover, larvae grow equally well on the foliage of damaged and undamaged pot-bound plants. The higher chlorophyll contents characteristic of damaged released plants did not negate the effects of the increased alkaloid contents on larval growth. Undamaged leaves from undamaged field-grown plants stem-fed nicotine solutions had elevated leaf nicotine and nornicotine contents. Larvae reared on these artificially induced leaves gain only 38.5% of the weight gained by larvae reared on low-alkaloid foliage. These results demonstrate that damage-induced increases in leaf alkaloids protect induced foliage from attack and are sufficient to explain the decreased growth of caterpillars on the foliage of damaged plants.     

Behavioral ecology ofMyiopharus doryphorae (riley) andM. aberrans (townsend), tachinid parasitoids of the colorado potato beetle

The behavior ofMyiopharus doryphorae andM. aberrans, North American tachinid parasitoids of the Colorado potato beetle, was recorded under field and laboratory conditions throughout three growing seasons in western Massachusetts. Eight common behaviors associated with resting, searching, feeding, and larviposition were distinguished, which together accounted for nearly all daytime activity of the females of both tachinids. Several of these behaviors, and in particular larviposition, were closely related to temperature but differed between species. A sequence of five defensive behaviors by the different larval stages of the Colorado potato beetle prevented larviposition in 49% of resisted attempts and perhaps one-fourth of total larviposition attempts byMyiopharus species, yet both parasitoids were highly successful in allocating their progeny during most of the summer. Second- and third-instar beetle larvae were least effective in resisting larviposition. Females of bothMyiopharus species actively guarded recently parasitized hosts from otherMyiopharus females for a period of several minutes after larviposition during the last month of the growing season when second- and third-instar Colorado potato beetle larvae were most scarce. Laboratory studies based on the field observation that femaleM. aberrans doggedly pursued circum-diapausing adult beetles led to the first recorded account ofM. aberrans larvipositing in adult hosts. Flies gained access to a beetle’s vulnerable abdominal dorsum at the instant it lifted its elytra to initiate flight. The late-season switch ofM. aberrans to adult Colorado potato beetles contributed to a seasonal sequence of larviposition-related behaviors concordant with prevailing host densities, which should lend complementarity toM. doryphorae andM. aberrans as biological controls of pest populations.     

Survival and fecundity of Bt-susceptible Colorado potato beetle adults after consumption of transgenic potato containing Bacillus thuringiensis subsp. tenebrionis Cry3A toxin

Survival and fecundity of Colorado potato beetle adults, Leptinotarsa decemlineata (Say), that had or had not fed previously on non-transgenic potato before exposure to transgenic potato containing the Bacillus thuringiensis subsp. tenebrionis Cry3A toxin (Bt) was investigated. In the laboratory, < 5% of first-generation adults survived after two weeks when restricted to Bt foliage since eclosion, but over 85% of adults that had fed initially on non-Bt potato survived exposure to Bt potato for two weeks. In field experiments, less than 0.5% of adults that were exclusively provided Bt potato plants survived overwinter, whereas 44% to 57% survived overwinter when fed non-Bt potato plants for two weeks before being provided Bt potato as a final pre-overwintering host. Survival through the winter increased as the duration of initial feeding on non-Bt potato increased and was similar for beetles provided either tubers or Bt potato plants as a final pre-overwintering host. Only overwintered beetles that fed initially on non-Bt potato before encountering either tubers or Bt potato as a final pre-overwintering host laid eggs the following spring. Survival and reproduction of potato beetle adults after colonizing Bt potato fields should not be adversely affected as long as they have had sufficient time to feed initially on non-Bt potato. Implications for how potato production practices in the Mid-Atlantic US may affect the utility of general resistance management plans for Bt potato are discussed.     

Systemic induction of volatile release in cotton: How specific is the signal to herbivory?

Plants attacked by herbivorous insects release chemical signals that attract natural enemies of the herbivores to the damaged plants. Feeding of Spodoptera exigua larvae on the lower leaves of cotton (Gossypium hirsutum L.) for multiple feeding periods of 9–12 h with a 12 h, interval in between when the caterpillars are removed overnight, will induce a systemic release of volatile compounds that is comparable to the volatiles released in response to continuous feeding damage on the lower leaves for several days. The systemic volatile release in response to herbivory can be mimicked by mechanically damaging the lower leaves and applying caterpillar oral secretion to the injured leaves over 4 days. Cotton plants that are only mechanically damaged systemically release significantly less -pinene, myrcene, (Z)-3-hexenyl acetate, (E)--farnesene and (E,E)--farnesene after 4 days compared to plants damaged mechanically with application of caterpillar regurgitant. However, multiple 9–12 h mechanical damage alone induces a significantly higher systemic release of (Z)-3-hexenyl acetate, myrcene, (E)--ocimene, and (E)--farnesene after 4 days compared to undamaged control plants. This indicates that multiple mechanical damage alone cannot mimic completely the response induced by mechanically injuring the leaves and applying caterpillar regurgitant. A specific elicitor in the regurgitant of the caterpillar enhances the amount of several systemically released volatiles. Thus, the systemic release of volatile compounds by herbivore-damaged cotton plants appears to be regulated by at least two different mechanisms.