Essential Compounds in Herbivore-Induced Plant Volatiles that Attract the Predatory Mite <Emphasis Type="Italic">Neoseiulus womersleyi</Emphasis>

Carnivorous arthropods use volatile infochemicals emitted from prey-infested plants in their foraging behavior. Although several volatile components are common among plant species, the compositions differ among prey–plant complexes. Studies showed that the predatory mite Neoseiulus womersleyi is attracted only to previously experienced plant volatiles. In this study, we identified the attractant components in prey-induced plant volatiles of two prey–plant complexes. N. womersleyi reared on Tetranychus kanzawai-infested tea leaves showed significant preference for a mixture of three synthetic compounds [mimics of the T. kanzawai-induced tea leaves volatiles: (E)-β-ocimene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), and (E,E)-α-farnesene] at a level comparable to that for T. kanzawai-induced tea plant volatiles. However, mixtures lacking any of these compounds did not attract the predatory mites. Likewise, N. womersleyi reared on T. urticae-infested kidney bean plants showed a significant preference for a mixture of four synthetic compounds [mimics of the T. urticae-induced kidney bean volatiles: DMNT, methyl salicylate (MeSA), β-caryophyllene, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene] at a level comparable to that for T. urticae-induced kidney bean volatiles. The absence of any of the four compounds resulted in no attraction. These results indicate that N. womersleyi can use at least four volatile components to identify prey-infested plants.     

Predator Avoidance by Phytophagous Mites is Affected by the Presence of Herbivores in a Neighboring Patch

When predators invade a leaf patch inhabited by herbivores, the herbivores disperse to a neighboring predator-free leaf patch, thus escaping from the predators. However, the neighboring patch might already be used by con- or heterospecific herbivores. We used laboratory bioassays to examine whether perception of odor from con- or heterospecific competitors on a neighbored lima bean leaf patch influences dispersal behavior of the herbivorous mite Tetranychus urticae when attacked by predatory mites Phytoseiulus persimilis. The dispersal rates of T. urticae that perceived odors from leaf patches infested by conspecifics or cowpea aphids (Aphis craccivora) did not differ from the control (the dispersal rate of T. urticae that perceived odor from uninfested leaf patches). By contrast, the dispersal rate of T. urticae was reduced when they perceived odors from leaf patches that were currently or had previously been infested by larvae of the common cutworm (Spodoptera litura). Previous herbivory by S. litura larvae induced resistance in leaf patches to T. urticae as indicated by the reduced number of eggs laid by T. urticae. Our results are discussed with respect to the feeding behavior of the tested competitors of T. urticae and the impact of the plant and arthropod community on the dispersal behavior of these mites.     

Contrasting Behavioral Responses by Detritivorous and Predatory Mayflies to Chemicals Released by Injured Conspecifics and Their Predators

Larvae of the mayfly Siphlonisca are predators of the detritivorous mayfly Siphlonurus in floodplain wetlands in Maine (USA). Both mayflies are natural prey of brook trout (Salvelinus fontinalis). We exposed larvae of Siphlonurus and Siphlonisca to chemicals from injured conspecifics and their predators. Significant decreases in movement activity by Siphlonurus were elicited by chemicals released from Siphlonisca, chemicals released from brook trout fed conspecifics, and by chemicals released from injured conspecifics. A significant decrease in movement activity by Siphlonisca was elicited by chemicals released from brook trout fed either conspecifics or Siphlonurus. Movement activity by either Siphlonurus or Siphlonisca was not significantly affected by chemicals released from trout feeding on brine shrimp (Artemia). Both Siphlonurus and Siphlonisca were able to detect chemicals that provided information about past feeding behaviour by brook trout. However, their response to the chemicals used in this study was context-specific. A reduction in movement activity, a behavior that presumably reduces the probability of being consumed by visual predators, occurred only when mayflies were exposed to chemicals released by brook trout feeding on conspecific (Siphlonurus) or confamilial (Siphlonisca) prey.     

Olfactory Responses of Neotropical Short-Tailed Singing Mice, <Emphasis Type="Italic">Scotinomys teguina</Emphasis>, to Odors of the Mid-Ventral Sebaceous Gland: Discrimination of Conspecifics,Gender, and Female Reproductive Condition

We examined the olfactory communication of the Neotropical short-tailed singing mouse, Scotinomys teguina, by investigating whether S. teguina responded to odors produced by the mid-ventral sebaceous gland of conspecifics. Females spent significantly more time investigating male odor than an odorless stimulus or a female odor. Males spent significantly more time investigating female odor than an odorless stimulus, but not that of a male odor. This latter result does not seem to be explained by differences in age or sexual experience of test subjects, but may be influenced by reproductive condition of the female odor donor. Male S. teguina spent significantly more time (1) investigating and (2) in total proximity to odors of estrous than non-estrous females. Males spent more time (1) investigating and (2) in total proximity to odors of males than non-estrous females. Finally, given the choice between odor of males vs proestrous females, males showed no preference. Thus, the mid-ventral gland in S. teguina seems to convey information about conspecifics, sex, and female reproductive condition. Male odor, compared with proestrous female odor, is equally interesting to males, suggesting that the gland also plays an important role in male–male communication. Sexual dimorphism in the size of the gland and in the amount of secretion produced by the gland may be related to either male–male competition or female choice.     

Influence of odor from wood-decaying fungi on host selection behavior of deathwatch beetle,Xestobium rufovillosum

Adult females of Xestobium rufovillosum de Geer demonstrated anemotactic orientation when exposed to an odor plume containing volatiles generated by wood-decaying fungi (Coriolus versicolor, Donkioporia expansa) and decayed oak wood (Quercus petraea, Quercus robur). They did not orient towards undecayed oak wood, beech (Fagus sylvatica), or pine wood (Pinus sylvestris). Although all insects tested showed anemotactic orientation, responses were nonlinear with respect to insect age. Adult females more readily oriented upwind when they were between 10 and 16 days old. Oviposition choice bioassays showed that ovipositing females would preferentially oviposit on extract-treated cellulose paper discs that had been treated with various strains of the wood-decaying fungus, Donkioporia expansa. HPLC-fractionated mycelial extracts were attractive to ovipositing deathwatch beetles, whereas HPLC-fractionated fungal broth extracts were repellent to ovipositing females. The implications of these results are discussed in the context of timber pest management in historically important buildings.     

Avoidance response of a terrestrial salamander (Ambystoma macrodactylum) to chemical alarm cues

Organisms from a wide variety of taxonomic groups possess chemical alarm cues that are important in mediating predator avoidance. However, little is known about the presence of such alarm cues in most amphibians, and in particular terrestrial salamanders. In this study we tested whether adult long-toed salamanders (Ambystoma macrodactylum) showed an avoidance response to stimuli from injured conspecifics. Avoidance of stimuli from injured conspecifics could represent avoidance of a chemical alarm cue or, alternatively, avoidance of a territorial pheromone or conspecific predator odor. Consequently, we also tested whether salamanders avoided stimuli from noninjured conspecifics. Salamanders avoided stimuli from injured but not from noninjured conspecifics. Therefore, we concluded that the response to injured conspecifics represents avoidance of a chemical alarm cue and not avoidance of a territorial pheromone or predator cue. This is the first clear demonstration of chemical alarm signaling by a terrestrial amphibian and the first report of chemical alarm signaling in an ambystomatid salamander. By avoiding an area containing stimuli from injured conspecifics, long-toed salamanders may lower their risk of predation by avoiding areas where predators are foraging.     

Exposure to odors from stressed conspecifics increases preference for higher ambient temperatures in C57BL/6J mice

Exposure to odors from stressed conspecifics increases preference for higher ambient temperatures in C57BL/6J mice. C57BL/6J male mice were individually allowed preferences on a thermal gradient ranging in temperature from 22°C to 42°C. Group 1 (N=10) was exposed to odors from triads of foot-shocked conspecifics during the first 2-hr temperature preference trial. Group 2 (N=10) was exposed to odors from triads of nonstressed conspecifics during similar testing. Body temperature (T_B) variations were measured in three animals of each group. Thermal preference was significantly higher for animals exposed to odors from stressed conspecifics than for animals exposed to odors from nonstressed animals (32.0°C vs. 29.0°C).T _B changes on the heated gradient were significantly higher for animals exposed to odors from stressed animals (+1.5°C) than for animals exposed to odors from nonstressed animals (–0.33°C). Additional animals on a non-heated thermogradient were tested for T_B when exposed to odors from stressed or from nonstressed animals (N=3 per condition). There was no difference inT _B between these two groups. Increases in T_B on the heated gradient are apparently due to the higher ambient temperature choices and not due to the odor per se.     

Potential use of chemical cues for colony-mate recognition in the big brown bat,Eptesicus fuscus

Bats should benefit from recognition of their roost-mates when colonies form stable social units that persist over time. We used Y-maze experiments and gas chromatography–olfactometry (GC-O) to evaluate whether female big brown bats Eptesicus fuscus (Chiroptera: Vespertilionidae) use chemical cues to distinguish among conspecifics. In dual-choice Y-maze experiments, females chose the scent of another female from their own roost over a conspecific female from a different roost in a majority of trials. Analysis of total body odors using GC-O suggests that individuals from a given colony may share a more common odor signature with roost-mates than with non-roost-mate conspecifics. Using four principle components derived from 15 odor variables, discriminant function analysis correctly assigned most individuals to the correct colony.     

Attraction and Oviposition of <Emphasis Type="Italic">Tuta absoluta</Emphasis> Females in Response to Tomato Leaf Volatiles

The tomato leafminer Tuta absoluta (Lepidoptera: Gelechiidae) is a devastating pest of cultivated tomato Solanum lycopersicum throughout South and Central America and Europe. We aimed to characterize the behavioral mechanisms and the chemical cues involved in host selection of T. absoluta females by chemical analysis of tomato leaf volatiles, wind tunnel attraction assays, and oviposition bioassays. Tomato leaf odor elicited in mated females upwind orientation flight followed by landing as well as egg-laying, demonstrating the essential role of plant volatiles in T. absoluta host-finding behavior. In wind tunnel and oviposition choice experiments, T. absoluta females significantly preferred tomato S. lycopersicum over wild tomato Solanum habrochaites, which is resistant to larval feeding. This indicates that leaf volatiles provide information on the suitability of plants as larval hosts. Mated females also discriminated three cultivars of S. lycopersicum according to their volatile profiles. Headspace collections from leaves of these three cultivars contained large amounts of β-phellandrene, followed by limonene, 2-carene, and (E)-β-caryophyllene, which together accounted for more than 70% of tomato foliage headspace. Most leaf volatiles were released by all three cultivars, but they showed significant differences with respect to the presence of a few minor compounds and blend proportion. This is an initial study of the volatile signatures that mediate attraction and oviposition of tomato leafminer T. absoluta in response to its main host, tomato.     

Odor Preferences of Weanling and Mature Male and Female Pine Voles

Olfactory cues from conspecifics have strong influences on behavior. Both female and male juveniles are attracted to familiar odors while adults show preferences for odors of opposite-sex conspecifics. Females show an increased preference for male odors as they mature, but no study has examined whether change with age is seen in males. In this experiment, responses of weanling and mature male and female pine voles (Microtus pinetorum) to familiar odor cues from their family versus those of an unfamiliar, opposite-sex conspecific were examined. Voles were tested in a three-chambered cage with end chambers containing the stimuli. There was a significant difference in odor preference between weanling and mature females. Weanling females preferred odor cues from their families, and mature females preferred the odors of unfamiliar mature males. The pattern was similar in males but the effect was not as strong. The results support the hypothesis that responses to odor cues change with age in pine voles. At least for females, the change in response to odor cues may be involved in the timing of reproduction and thus dispersal.     

Identification of male cabbage looper sex pheromone attractive to females

A multicomponent pheromone produced by male cabbage looper moths that is attractive to female moths in a flight tunnel bioassay was isolated and identified. Based on analyses of hairpencil extracts of male cabbage loopers and volatiles emitted by males, the pheromone has been identified as a blend consisting of (S)-(+)-linalool,p-cresol, andm-cresol. The chirality of the major component, (S)-(+)-linalool, is important for behavioral response of females. These pheromonal compounds were also identified as volatiles released by males when males were exposed to the principal pheromone component of female cabbage loopers, (Z)-7-dodecen-1-ol acetate. The amount of male pheromone released was increased significantly when males were exposed to a combination of (Z)-7-dodecen-1-ol acetate and the odor from cabbage. Neither linalool nor the cresols were detected in volatiles from cabbage or from males exposed to cabbage odor.     

Alert odor from skin gland in deer

Black-tailed deer (Odocoileus hemionus columbianus) discharge an odor from the metatarsal (MT) gland, located on the hindleg, when disturbed or alarmed. Freely moving, captive deer were exposed to the MT odor by means of a remote-release apparatus. Responses by males and females to MT odor from both sexes were recorded with a coding system and a video camera. When the odor was present, females became more alert and left the site more often than in the presence of control odors, odorless air, or deer urine. It is concluded that the MT secretion provides an alert signal, placing the odor into the class of alarm pheromones. There is no evidence that the closely related white-tailed deer,O. virginianus, possesses this alert (or alarm) odor system to the same degree.     

Perception of Conspecific Female Pheromone Stimulates Female Calling in an Arctiid Moth, <Emphasis Type="Italic">Utetheisa ornatrix</Emphasis>

Perception of the female sex pheromone in Utetheisa ornatrix (Lepidoptera: Arctiidae) is responsible for induction and adjustment of calling by females and the collective phenomenon termed “female pheromonal chorusing”. We found five olfactory-active compounds in the U. ornatrix female gland. When females were exposed to the entire pheromone or to two of its (synthetically prepared) components, (Z,Z,Z)-3,6,9-eicosatriene and (Z,Z,Z)-3,6,9-heneicosatriene, they were more likely to call during a given night, begin calling earlier, and briefly increase signal frequency with which they extrude their abdomen, an observable indication of calling in this species. Some females even initiated calling during photophase when exposed to the pheromone components. In general, female U. ornatrix are more sensitive to the complete blend of pheromone than to its individual compounds. We also tested the hypotheses: 1) that abdominal extrusion per se increases the rate of pheromone release; and 2) that greater abdominal pumping rhythm increases pheromone release rate. Contrary to our expectations: 1) females did not respond more strongly to a pulsed pheromone stimulus than to the constant release of pheromone at the same average release rate; and 2) we did not find a relationship between the frequency of abdominal pumping and pheromone release rate. Possible explanations for these unexpected findings are discussed.     

Flight Initiation by Male <Emphasis Type="Italic">Rhodnius prolixus</Emphasis> is Promoted by Female Odors

Several triatomine bug species utilize chemical cues for sexual communication. We tested whether female or male Rhodnius prolixus, a vector of Chagas disease, produce volatile chemicals that elicit flight responses from conspecifics, and then isolated the source of the chemical. Males confronted with an airstream containing female odors showed a significantly greater take-off frequency compared to a blank airstream or an airstream with male odors. In contrast, females exhibited similar take-off frequencies to male or female odor as to a clean airstream. Occlusion of female metasternal glands with paraffin wax resulted in a significant decrease in male take-off frequency compared to that of intact females. Additionally, excised female metasternal glands elicited a similar take-off frequency from males as did intact females, both significantly greater than the take-off frequency to clean air. These results show that R. prolixus females release a pheromone from their metasternal glands that causes upwind flight in conspecific males.     

Parasitoids Modify Their Oviposition Behavior According to the Sexual Origin of Conspecific Cuticular Hydrocarbon Traces

Hydrocarbons play a crucial role in insect behavior in general and in sexual recognition in particular. Parasitoids often modify their oviposition behavior according to hydrocarbons left by conspecifics on the reproductive patch, such as oviposition markers left by females after oviposition, or cuticular hydrocarbon (CHC) traces left by individuals by walking or rubbing. This study determined whether Eupelmus vuilleti females are able to distinguish CHCs left by male or female conspecifics on seeds. The results show that the cuticular profile of E. vuilleti differs according to its gender, and that females are able to detect the sexual origin of these CHCs. Moreover, they adjust their oviposition behavior according to the nature of these traces. Although females lay fewer eggs on hosts when confronted with female CHCs, they lay more daughters when confronted with male CHCs, thus changing the sex ratio.     

Antiaphrodisiacs in pierid butterflies: a theme with variation!

Male Pieris napi butterflies previously have been shown to synthesize and transfer an antiaphrodisiac, methyl salicylate (MeS), to females at mating. This substance curtails courtship and decreases the likelihood of female remating. Here, we show that similar systems occur in Pieris rapae and Pieris brassicae. In P. rapae, ¹³C-labeling studies showed that males utilize the amino acids phenylalanine and tryptophan as precursors to MeS and indole, respectively. These volatiles are transferred to females at mating and function as antiaphrodisiacs, as demonstrated by field tests entailing painting MeS, indole, or a mixture on the abdomens of virgin females and assessing their attractiveness to wild males. With P. brassicae, ¹³C-labeling studies showed that males use phenylalanine as a precursor to synthesize benzyl cyanide, which was demonstrated to function as an antiaphrodisiac by field tests similar to those for P. rapae. This communication system exhibits both similarities and differences among the three species; in P. napi and P. rapae, males are fragrant but transfer a volatile antiaphrodisiac to females that is completely different from the male odor, whereas in P. brassicae the antiaphrodisiac transferred by male to female is identical with male odor.     

Effect of predator diet on life history shifts of red-legged frogs,Rana aurora

Larval red-legged frogs (Rana aurora) are known to exhibit antipredator behavior in response to both chemical alarm cues released from injured conspecifics and chemical cues of predators. In some cases, the response to predators is dependent on the predator's diet. In this experiment, we tested whether long-term exposure to predator chemical cues and alarm cues resulted in alteration of life history characteristics of red-legged frogs. We raised groups of tadpoles in the presence of chemical cues of predators that were either fed conspecifics or heterospecific caddis-fly larvae, chemical cues of injured conspecifics, or a no-cue control. Tadpoles raised in the presence of either a predator fed conspecifics or cues of injured conspecifics metamorphosed earlier and at a smaller size than those exposed to predators fed heterospecifics or the no-cue control. The result suggests that red-legged frogs exhibit a life history shift in response to predatory cues and that this response is dependent on the diet of predators.     

Intraspecies Variation in the Kanzawa Spider Mite Differentially Affects Induced Defensive Response in Lima Bean Plants

The Kanzawa spider mite, Tetranychus kanzawai, is a polyphagous herbivore that feeds on various plant families, including the Leguminacae. Scars made by the mite on lima bean leaves (Phaseolus lunatus) were classified into two types: white and red. We obtained two strains of mites—“White” and “Red”—by selecting individual mites based on the color of the scars. Damage made by the Red strain induced the expression of genes for both basic chitinase, which was downstream of the jasmonic acid (JA) signaling pathway, and acidic chitinase, which was downstream of the salicylic acid (SA) signaling pathway. White strain mites also induced the expression of the basic chitinase gene in infested leaves but they only slightly induced the acidic chitinase gene. The Red genotype was dominant over the White for the induction of the acidic chitinase gene. The amount of endogenous salicylates in leaves increased significantly when infested by Red strain mites but did not increase when infested by White strain mites. JA and SA are known to be involved in the production of lima bean leaf volatiles induced by T. urticae. The blend of volatiles emitted from leaves infested by the Red strain were qualitatively different from those infested by the White strain, suggesting that the SA and JA signaling pathways are differently involved in the production of lima bean leaf volatiles induced by T. kanzawai of different strains.Ryo Matsushima and Rika Ozawa contributed equally to this work.     

Behavioral responses ofLittoraria irrorata (SAY) to water-borne odors

Behavioral responses of the gastropod molluscLittoraria (=Littorina)irrorata indicate that it can discriminate among environmental odors. Snails were assayed for responses to 11 odors from plants and animals potentially representing food, shelter, location in the environment, and predators. Crushed conspecifics were included as an alarm odor. Except for odor of crushed conspecifics, all odor sources were water-borne from living intact organisms. Behavioral responses were categorized as no response, positive response, or negative response. For some analyses, negative responses were subdivided into withdrawing and turning responses. Snails responded positively to several plant odors. They did not respond to odors of intact conspecifics, fiddler crabs, or grass shrimp. They responded negatively to odors of a plant found at the upper limit of their minimal habitat, predatory blue crabs, crushed conspecifics, predatory gastropods, and ribbed mussels. Odors of blue crabs on different diets affect the type of negative response the snails display.     

Behavioral responses of sexually active mud snails: kariomones and pheromones

We postulated that kariomones and pheromones function to organize egg capsule deposition and breeding aggregations of mud snails (Ilyanassa obsoleta). Sexually active male and female mud snails were exposed to water-born odors from juvenile snails, nonaggregated female snails, oysters, egg capsule depositing snails, sexually active males, sexually active females, and copulating snails and tested for a follow response using a ring assay. Neither sex responded to juvenile odor or to odor of sexually inactive females. Snails turned and followed water born odors from oysters. when tested with the odors of sexually active males, sexually active females, and copulating pairs, we found sex-specific differences in responses. Sexually active females followed the odor of sexually active males and ignored sexually active female odor. Sexually active males followed the odor of sexually active females and ignored sexually active male odor. Both sexes followed the odor of copulating pairs and egg capsule depositing snails. Snail activities in breeding aggregations are organized by at least three pheromones and one kariomone, and reflect the multifunctionaLity of the aggregations.