Role of olfaction in host finding by aphid parasitoidAphidius nigripes (Hymenoptera: Aphidiidae)

The olfactory responses of the parasitoidAphidius nigripes Ashmead were investigated with a four-field airflow olfactometer. Female and male parasitoids did not respond to potato plants, the food plant of their host. However, females were strongly attracted by odors from a preferred host,Macrosiphum euphorbiae (Thomas), and also from less preferred hosts such asMyzus persicae (Sulzer) andAphis nasturtii Kalt. Moreover,A. nigripes females responded positively to odors from a nonhost aphid,Rhopalosiphum maidis Fitch. The honeydew produced by these four aphid species was also very attractive to females. Males did not respond to aphids or honeydew but were highly attracted by odors from conspecific female parasitoids.Contribution No. 394 from the Département de biologie, Université Laval.     

Olfactory Responses of Parasitoid Apoanagyrus lopezi to Odor of Plants, Mealybugs, and Plant–Mealybug Complexes

Apoanagyrus (Epidinocarsis) lopezi De Santis is an endoparasitoid used in the biological control of the cassava mealybug Phenacoccus manihoti Matile-Ferrero in Africa. The response of naive and mated females of A. lopezi to odors from cassava plant (var. Zanaga), parasitized or unparasitized mealybugs, and plant–mealybug host complexes with or without feeding hosts was investigated in a Y-tube olfactometer. Dual-choice tests revealed that mealybug-infested plants and mealybug-damaged plants were the major sources of volatiles that attract female parasitoids to the microhabitat of its hosts. The emission of volatile chemicals appears not to be limited to the infested plant part but to occur systemically throughout the plant. On their own, unparasitized mealybugs were more attractive than uninfested plants or parasitized mealybugs alone. Parasitization of P. manihoti by A. lopezi decreased the response of parasitoids to mealybugs or mealybug–plant complexes. Plants infested with unparasitized hosts attracted more female parasitoids than plants infested with parasitized mealybugs. These results indicate that, in the long-range host-searching process, females of A. lopezi respond mainly to mealybug-induced synomones, and specific host-derived cues play a minor role.     

Olfactory responses of the parasitoidDiaeretiella rapae (Hymenoptera: Aphidiidae) to odor of plants,aphids, and plant-aphid complexes

Diaeretiella rapae (M'Intosh) (Hymenoptera: Aphidiidae) is a parasitoid of several aphid species, including the Russian wheat aphid (RWA),Diuraphis noxia (Mordvilko), and the cabbage aphid (CA).Brevicoryne brassicae (L.). The response of matedD. rapae females to odors from wheat, cabbage, and plant-host complexes was investigated using a four-choice olfactometer. Experienced parasitoids, but not inexperienced females, responded positively to odors of the wheat-RWA complex in a no-choice test. In choice tests, experienced parasitoids did not respond to odors of uninfested cabbage and wheat leaves, but did respond positively to aphid-infested plants and to aphids alone. The response ofD. rapae to the cabbage-CA complex and to CA alone was significantly greater than to the wheat-RWA complex and RWA alone, suggesting an innate odor preference for crucifer-feeding aphids.     

Odor Perception in the Bark Beetle Parasitoid Roptrocerus xylophagorum Exposed to Host Associated Volatiles

Y-tube olfactometer bioassays and combined gas chromatography–electroantennographic detector (GC-EAD) analyses were performed to investigate the specific odors utilized as host location cues by the bark beetle parasitoid, Roptrocerus xylophagorum, originating from the southeastern United States. R. xylophagorum parasitizes several economically important holarctic bark beetle species and females oviposit preferentially on late larval stages. Both female and male parasitoids were tested with volatiles derived from host infestations of either Dendroctonus frontalis, the southern pine beetle, or Ips grandicollis, the southern pine engraver. Tested volatiles were steam distillates from the bark of loblolly pine, Pinus taeda, infested with larvae of the respective bark beetle species. Combined gas chromatography–mass spectrometry (GC-MS) was employed for identification and quantification of the compounds in the steam distillates. To confirm the EAD activity of identified compounds, GC-EAD analyses were repeated with a synthetic blend composed predominantly of compounds in the crude extracts that had revealed apparent electrophysiological activity. In Y-olfactometer bioassays, female parasitoids were attracted to both of the above-mentioned distillates. Male parasitoids were tested with one of the distillates but failed to respond. In GC-EAD analyses, the sexes displayed similar relative sensitivities to the components of the blends. Males exhibited generally higher amplitudes of response to the tested compounds than females. Monoterpene hydrocarbons associated with the constitutive resin of the host tree did not elicit significant EAD responses. Compounds known to be associated specifically with the host–tree complex, such as certain oxygenated monoterpenes, generated the greatest EAD responses. Female parasitoids were attracted by a synthetic blend composed of several of the EAD active oxygenated monoterpenes.     

Differential attractiveness of induced odors emitted by eight maize varieties for the parasitoid cotesia marginiventris: is quality or quantity important?

Herbivore-induced plant volatiles can function as indirect defense signals that attract natural enemies of herbivores. Several parasitoids are known to exploit these plant-provided cues to locate their hosts. One such parasitoid is the generalist Cotesia marginiventris, which is, among others, attracted to maize volatiles induced by caterpillar damage. Maize plants can be induced to produce the same blend of attractive volatiles by treating them with regurgitant of Spodoptera species. We collected and analyzed the regurgitant-induced emissions of two plant species (cowpea and maize) and of eight Mexican maize varieties and found significant differences among their volatile emissions, both in terms of total quantity and the quality of the blends. In a Y-tube olfactometer, the odors of the same artificially induced plant species and Mexican varieties were offered in dual choice experiments to naïve mated females of C. marginiventris. Wasps preferred cowpea over maize odor and, in 3 of 12 combinations with the maize varieties, they showed a preference for the odors of one of the varieties. A comparison of the odor collection with results from the behavioral assays indicates that not only the quantity of the volatile emissions, but also the quality composition of the volatile blends is important for attraction of C. marginiventris. The results are discussed in the context of the possibility of breeding crop varieties that are particularly attractive to parasitoids.     

Effect of the Presence of a Nonhost Herbivore on the Response of the Aphid Parasitoid <Emphasis Type="Italic">Diaeretiella rapae</Emphasis> to Host-infested Cabbage Plants

The vast majority of studies of plant indirect defense strategies have considered simple tritrophic systems that involve plant responses to attack by a single herbivore species. However, responses by predators and parasitoids to specific, herbivore-induced, volatile blends could be compromised when two or more different herbivores are feeding on the same plant. In Y-tube olfactometer studies, we investigated the responses of an aphid parasitoid, Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae), to odors from cabbage plants infested with the peach-potato aphid Myzus persicae (Sulzer) (Homoptera: Aphididae), in both the presence and absence of a lepidopteran caterpillar, Plutella xylostella L. (Lepidoptera: Plutellidae). Female parasitoids chose aphid-infested plants over uninfested plants but did not distinguish between caterpillar-infested and uninfested plants. When given a choice between odors from an aphid-infested plant and those from a plant infested with diamondback moth larvae, they significantly chose the former. Furthermore, the parasitoids responded equally to odors from a plant infested with aphids only and those from a plant infested with both aphids and caterpillars. The results support the hypothesis that the aphid and the caterpillar induce different changes in the volatile profile of cabbage plants and that D. rapae females readily distinguish between the two. Furthermore, the changes to the plant volatile profile induced by the caterpillar damage did not hinder the responses of the parasitoid to aphid-induced signals.     

Volatile infochemicals used in host and host habitat location byCotesia flavipes Cameron andCotesia sesamiae (Cameron) (Hymenoptera: Braconidae), larval parasitoids of stemborers on graminae

FemaleCotesia flavipes Cameron andCotesia sesamiae (Cameron) were attracted to odors in a Y-tube olfactometer from uninfested maize (Zea mays L.), sorghum [Sorghum bicolor (L.)], and napier grass (Pennisetum purpureum Schumach). In dual choice tests,Cotesia flavipes showed a preference for maize over sorghum, while maize and napier grass were equally attractive. In contrast,Cotesia sesamiae preferred volatiles from sorghum and napier grass over those from maize. The two parasitoids were significantly more attracted to maize infested with the stemborers,Chilo partellus (Swinhoe),Chilo orichalcociliellus Strand,Sesamia calamistis Hampson, andBusseola fusca (Fuller), than uninfested maize. In dual choice tests,Cotesia flavipes andCotesia sesamiae were unable to discriminate between odors from plants infested by the different species of stemborers.     

Semiochemicals from Fungal Associates of Bark Beetles May Mediate Host Location Behavior of Parasitoids

In laboratory olfactometer bioassays, females of two hymenopteran parasitoid species, Roptrocerus xylophagorum (Ratzeburg) (Hymenoptera: Pteromalidae) and Spathius pallidus (Ashmead) (Hymenoptera: Braconidae), were attracted to odors from bark or bolts of loblolly pine, Pinus taeda L., colonized by bluestain fungi (genus Ophiostoma) associated with the parasitoids' bark beetle hosts. Mock-inoculated bolts and bark were less attractive or unattractive in these bioassays. Bark infested with host larvae that lacked their fungal and other normal microbial associates was attractive to R. xylophagorum females, but was less so than bark infested with larvae possessing their normal complement of associated microbes. In contrast, in oviposition bioassays, R. xylophagorum females spent approximately equal time searching, made similar numbers of oviposition attempts, parasitized similar percentages of hosts, and laid similar numbers of eggs in bark fragments infested with either associate-free or associate-bearing host larvae. Furthermore, in field bioassays using bluestain-inoculated or mock-inoculated loblolly pine bolts as sources of attractants, the numbers of parasitoids attracted by the two treatments did not differ significantly and the two treatments were less attractive than bolts naturally infested with bark beetle larvae. Whereas our laboratory olfactometer data suggest that bark beetle fungal associates may enhance attraction of some parasitoids, our bioassays with associate-free hosts indicate that associate-produced are not required for short-range host location and parasitization. In addition, our field trials indicated that long-range attraction of parasitoids to the host-fungi-tree complex is not caused simply by an interaction between bluestain fungi and tree tissues.     

Sex Attractant Pheromone from the Neotropical Red-Shouldered Stink Bug, <Emphasis Type="Italic">Thyanta perditor</Emphasis> (F.)

Olfactometer bioassays showed that odors from mature Thyanta perditor males attracted females but not males. Furthermore, odors from females did not attract either sex, indicating that like other phytophagous pentatomid bugs, the males produce a sex pheromone. Attraction appeared to peak in late afternoon to evening. The headspace volatiles collected from male and female T. perditor were analyzed by GC-MS and HPLC. A male-specific compound, methyl (2E,4Z,6Z)-decatrienoate (2E,4Z,6Z-10:COOMe), was identified along with a number of other compounds found in extracts from both sexes. Bioassays carried out with 2E,4Z,6Z-10:COOMe showed it was as attractive to females as the crude extract of male volatiles, suggesting that the male-produced sex pheromone consists of 2E,4Z,6Z-10:COOMe as a single component. Consecutive volatiles collections from males showed that 2E,4Z,6Z-10:COOMe began appearing in extracts from males about 9 d after the final molt, as the males became sexually mature.     

Herbivore-Induced Plant Volatiles Mediate In-Flight Host Discrimination by Parasitoids

Herbivore feeding induces plants to emit volatiles that are detectable and reliable cues for foraging parasitoids, which allows them to perform oriented host searching. We investigated whether these plant volatiles play a role in avoiding parasitoid competition by discriminating parasitized from unparasitized hosts in flight. In a wind tunnel set-up, we used mechanically damaged plants treated with regurgitant containing elicitors to simulate and standardize herbivore feeding. The solitary parasitoid Cotesia rubecula discriminated among volatile blends from Brussels sprouts plants treated with regurgitant of unparasitized Pieris rapae or P. brassicae caterpillars over blends emitted by plants treated with regurgitant of parasitized caterpillars. The gregarious Cotesia glomerata discriminated between volatiles induced by regurgitant from parasitized and unparasitized caterpillars of its major host species, P. brassicae. Gas chromatography-mass spectrometry analysis of headspace odors revealed that cabbage plants treated with regurgitant of parasitized P. brassicae caterpillars emitted lower amounts of volatiles than plants treated with unparasitized caterpillars. We demonstrate (1) that parasitoids can detect, in flight, whether their hosts contain competitors, and (2) that plants reduce the production of specific herbivore-induced volatiles after a successful recruitment of their bodyguards. As the induced volatiles bear biosynthetic and ecological costs to plants, downregulation of their production has adaptive value. These findings add a new level of intricacy to plant–parasitoid interactions.     

Sesamia calamistis calling behavior and its role in host finding of egg parasitoids Telenomus busseolae,telenomus isis,and Lathromeris ovicida

In the Republic of Benin, the African pink stem borer Sesamia calamistis Hampson is kept under control by the two scelionid egg parasitoids Telenomus busseolae (Gahan) and T. isis (Polaszek). A third species, the trichogrammatid Lathromeris ovicida Risbec, although commonly obtained from egg batches, is of low importance. High egg and egg batch parasitism by Telenomus spp. at low host densities suggests excellent host-finding capacity. Previously it was hypothesized that pheromones produced by calling S. calamistis virgins play an important role in attracting the parasitoids to the area where oviposition is likely to occur. Thus, after determining the effect of age (1–4 days) on the calling rhythm of S. calamistis, 1-day-old virgin females were chosen to investigate the response of the parasitoids to calling and noncalling females and two empty controls, in a four-arm olfactometer. The results showed that calling started earliest and lasted longest with 1-day-old females, but was similar for 2- to 4-days-old females. The percentage of females calling was higher in early than latter scotophases. The highest percentage of active females was obtained between 8 and 9 hr after the beginning of the first scotophase. All three parasitoid species responded to calling females, while there were no differences between noncalling females and the empty controls. The response pattern was similar for the two Telenomus spp. but considerably weaker for L. ovicida, reflecting differing host specificity of the three parasitoid species.     

Induction of volatile emissions in maize by different larval instars of Spodoptera littoralis

Maize plants under attack by caterpillars emit a specific blend of volatiles that is highly attractive to parasitic wasps. The release of these signals is induced by elicitors in the caterpillar regurgitant. Studies suggest that plants respond differently to different herbivore species and even to different herbivore stages, thus providing parasitoids and predators with specific signals. We tested if this is the case for different larval instars of the noctuid moth Spodoptera littoralis when they feed on maize plants. Cut maize plants were incubated in diluted regurgitant from second, third, or fifth instar caterpillars. There were no differences in total amount released after these treatments, but there were small differences in the release of the minor compounds phenethyl acetate and -humulene. Regurgitant of all three instars contained the elicitor volicitin. To test the effect of actual feeding by the larvae, potted plants were infested with caterpillars of one of the three instars, and volatiles were collected the following day. The intensity of the emissions was correlated with the number of larvae feeding on a plant, and with the amount of damage inflicted, but was independent of the instar that caused the damage. We also used artificial damage to mimic the manner of feeding of each instar to test the importance of physical aspects of damages for the odor emission. The emission was highly variable, but no differences were found among the different types of damage. In olfactometer tests, Microplitis rufiventris, a parasitoid that can only successfully parasitize second and early third instar S. littoralis, did not differentiate among the odors of maize plants attacked by different instar larvae. The odor analyses as well as the parasitoid's responses indicate that maize odors induced by S. littoralis provide parasitoids with poor information on the larval developmental stage. We discuss the results in the context of variability and lack of specificity in odorous plant signals.     

The Role of Fresh <Emphasis Type="Italic">versus</Emphasis> Old Leaf Damage in the Attraction of Parasitic Wasps to Herbivore-Induced Maize Volatiles

The odor produced by a plant under herbivore attack is often used by parasitic wasps to locate hosts. Any type of surface damage commonly causes plant leaves to release so-called green leaf volatiles, whereas blends of inducible compounds are more specific for herbivore attack and can vary considerably among plant genotypes. We compared the responses of naïve and experienced parasitoids of the species Cotesia marginiventris and Microplitis rufiventris to volatiles from maize leaves with fresh damage (mainly green leaf volatiles) vs. old damage (mainly terpenoids) in a six-arm olfactometer. These braconid wasps are both solitary endoparasitoids of lepidopteran larvae, but differ in geographical origin and host range. In choice experiments with odor blends from maize plants with fresh damage vs. blends from plants with old damage, inexperienced C. marginiventris showed a preference for the volatiles from freshly damaged leaves. No such preference was observed for inexperienced M. rufiventris. After an oviposition experience in hosts feeding on maize plants, C. marginiventris females were more attracted by a mixture of volatiles from fresh and old damage. Apparently, C. marginiventris has an innate preference for the odor of freshly damaged leaves, and this preference shifts in favor of a blend containing a mixture of green leaf volatiles plus terpenoids, after experiencing the latter blend in association with hosts. M. rufiventris responded poorly after experience and preferred fresh damage odors. Possibly, after associative learning, this species uses cues that are more directly related with the host presence, such as volatiles from host feces, which were not present in the odor sources offered in the olfactometer. The results demonstrate the complexity of the use of plant volatiles by parasitoids and show that different parasitoid species have evolved different strategies to exploit these signals.     

Relative importance of semiochemicals from first and second trophic levels in host foraging behavior ofAphidius ervi

The responses of femaleAphidius ervi to odors from a host food plant (Vicia faba), host aphids (Acyrthosiphon pisum), nonhost aphids (Aphis fabae), and aphid-plant complexes were investigated in a specially designed wind tunnel and a Y-tube olfactometer. In single-target (no-choice) and two-target (dual-choice) experiments, plant volatiles played a crucial role in the host foraging behavior ofA. ervi. The odor from theA. pisum-plant complex elicited the strongest responses byA. ervi females, followed by the odor from plants previously damaged by the feeding ofA. pisum. There was a significantly weaker response to odor fromA. pisum in the absence of the plant and to undamaged plants. Similarly, mechanically damaged plants and plants infested with the nonhost aphidA. fabae did not elicit strong responses. A plant that had been damaged byA. pisum and subsequently washed with distilled water was as attractive as an unwashed, previously infested plant.Aphidius ervi probably overcomes the reliability-detectability problem by selectively responding to herbivore-induced, volatile, semiochemical cues emitted by the first trophic level and by distinguishing between the volatiles induced by host and nonhost aphids.     

Behavioral and electrophysiological responses of the banana weevilCosmopolites sordidus to host plant volatiles

Male and femaleCosmopolites sordidus were attracted to freshly cut banana rhizome and pseudostem in a still-air olfactometer. Females responded similarly to odors from a comparatively resistant and from a susceptible cultivar of banana, when presented as either freshly cut tissue or as Porapak-trapped volatiles. Females were also attracted to rotting banana pseudostem and to volatiles collected from it. Males and females gave similar responses to host tissue in both the behavioral bioassay and to collected volatiles in EAG recordings. Weevils did not respond, either behaviorally or electrophysiologically, to a synthetic mixture of mono- and sesqiterpenes, which made up over 9% of the volatiles collected from pseudostem.     

Specific foraging kairomones used by a generalist parasitoid

In general, it is assumed that generalist natural enemies do not innately use specific cues for the location of their host or prey species. This hypothesis was tested using naïve females of the generalist parasitoid Lariophagus distinguendus Förster and two of its hosts, larvae of the lesser grain borer Rhyzopertha dominica (F.) and of the granary weevil Sitophilus granarius L., feeding in wheat grains. In a four-chamber olfactometer, female parasitoids were attracted to volatiles emanating from the feces of both host species. Chemical analysis of the volatiles from the feces of R. dominica revealed the presence of dominicalure 1 and 2, the species specific aggregation pheromones of R. dominica. The main compounds in the volatiles from feces of S. granarius were identified as chemicals related to mites that are associated with hosts of L. distinguendus. Because these mites are not specific for S. granarius but also co-occur with other hosts, the mite chemicals have to be considered as general cues. In bioassays, synthetic dominicalure was attractive to naïve L. distinguendus, explaining the attraction of feces volatiles from R. dominica. Synthetic mite chemicals and sitophilate, the aggregation pheromone of S. granarius, had no effect on naïve parasitoids. It remains to be determined which innate chemical cues from feces of S. granarius are used by L. distinguendus. In contrast to our initial hypothesis, the generalist L. distinguendus is innately using specific cues for foraging. Two ideas are provided to explain this result.     

Systemic release of herbivore-induced plant volatiles by turnips infested by concealed root-feeding larvae Delia radicum L

When attacked by herbivorous insects, many plants emit volatile compounds that are used as cues by predators and parasitoids foraging for prey or hosts. While such interactions have been demonstrated in several host–plant complexes, in most studies, the herbivores involved are leaf-feeding arthropods. We studied the long-range plant volatiles involved in host location in a system based on a very different interaction since the herbivore is a fly whose larvae feed on the roots of cole plants in the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The parasitoid studied is Trybliographa rapae Westwood (Hymenoptera: Figitidae), a specialist larval endoparasitoid of D. radicum. Using a four-arm olfactometer, the attraction of naive T. rapae females toward uninfested and infested turnip plants was investigated. T. rapae females were not attracted to volatiles emanating from uninfested plants, whether presented as whole plants, roots, or leaves. In contrast, they were highly attracted to volatiles emitted by roots infested with D. radicum larvae, by undamaged parts of infested roots, and by undamaged leaves of infested plants. The production of parasitoid-attracting volatiles appeared to be systemic in this particular tritrophic system. The possible factors triggering this volatile emission were also investigated. Volatiles from leaves of water-stressed plants and artificially damaged plants were not attractive to T. rapae females, while volatiles emitted by leaves of artificially damaged plants treated with crushed D. radicum larvae were highly attractive. However, T. rapae females were not attracted to volatiles emitted by artificially damaged plants treated only with crushed salivary glands from D. radicum larvae. These results demonstrate the systemic production of herbivore-induced volatiles in this host-plant complex. Although the emission of parasitoid attracting volatiles is induced by factors present in the herbivorous host, their exact origin remains unclear. The probable nature of the volatiles involved and the possible origin of the elicitor of volatiles release are discussed.     

Beneficial arthropod behavior mediated by airborne semiochemicals. II. Olfactometric studies of host location by the parasitoidMicroplitis croceipes (Cresson) (Hymenoptera: Braconidae)

The response ofMicroplitis croceipes (Cresson) to odors from a plant-host complex was investigated using a four-choice olfactometer. Female, but not male, parasitoids responded in a dose-dependent fashion to volatiles from the plant-host complex and oviposition experience enhanced this response. Female age had no apparent effect on the response. Both artificially damaged leaves and frass elicited positive responses but of lower magnitude than those elicited by the plant-host complex. Volatiles collected from the plant-host complex placed on filter paper also elicited positive responses by female parasitoids.Mention of a commercial or proprietary product does not constitute an endorsement by the USDA.     

Evidence of an aggregation pheromone in the flea beetle,Phyllotreta Cruciferae (Goeze) (Coleoptera: Chrysomelidae)

Laboratory olfactometer bioassays and field trapping experiments showed that the flea beetle,Phyllotreta cruciferae (Goeze), was highly attracted by oilseed rape(Brassica napus L.) when flea beetles were on the plant. This attraction was mediated by a flea beetle-produced aggregation pheromone based upon: (1) Oilseed rape damaged mechanically, or byP. cruciferae, or by diamondback moth,Plutella xylostella (L.), did not attractP. cruciferae. (2) Contact with the plants or feeding was required for the production of aggregation pheromone because oilseed rape alone was not attractive when separated from flea beetles by a screen. (3) Equal numbers of males and females were attracted.     

Plant–herbivore–carnivore Interactions in Cotton, <Emphasis Type="Italic">Gossypium hirsutum</Emphasis>: Linking Belowground and Aboveground

Most studies on plant–herbivore interactions focus on either root or shoot herbivory in isolation, but above- and belowground herbivores may interact on a shared host plant. Cotton (Gossypium spp.) produces gossypol and a variety of other gossypol-like terpenoids that exhibit toxicity to a wide range of herbivores and pathogens. Cotton plants also can emit herbivore-induced volatile compounds at the site of damage and systemically on all tissues above the site of damage. As these volatile compounds attract natural enemy species of the herbivore, they are thought to represent an indirect plant defense. Our study quantified gossypol and gossypol-like compounds in cotton plants with foliage feeding (Heliocoverpa zea), root feeding (Meloidogyne incognita), or their combination. Cotton plants with these treatments were studied also with respect to induced local and systemic volatile production and the attraction of the parasitic wasp Microplitis croceipes to those plants. We also evaluated whether foliage or root feeding affected foliar nitrogen levels in cotton. After 48 hr of leaf feeding and 5 wk of root feeding, local and systemic induction of volatiles (known to attract parasitoids such as M. croceipes) occurred with herbivore damage to leaves, and it increased in levels when root herbivory was added. Nevertheless, M. croceipes were equally attracted to plants with both leaf and root damage and leaf damage only. In contrast to previous studies in cotton, production of gossypol and gossypol-like compounds was not induced in leaf and root tissue following foliage or root herbivory, or their combination. We conclude that root feeding by M. incognita has little influence on direct and indirect defenses of Gossypium hirsutum against insect herbivory.