Ovipositional Behavior of Lesser Peachtree Borer in Presence of Host-Plant Volatiles 1
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Journal of Chemical Ecology, Vol. 14, No.1, 1988 OVIPOSITIONAL BEHAVIOR OF LESSER PEACHTREE BORER IN PRESENCE OF HOST-PLANT VOLATILES 1 D.K. REED,2,3 K.L. MIKOLAJCZAK,4 and C.R..KRAUSE5 3Fruit and Vegetable Insects Research Laboratory ARS, USDA, Vincennes, Indiana 47591 4Northern Region Research Center ARS, USDA, Peoria, Illinois 61604 5Nursery Crops Research Laboratory ARS, USDA, Delaware, Ohio 43015 (Received September 24, 1986; accepted January 22, 1987) Abstract-Reactions of lesser peachtree borer [Synanthedon pictipes (G&R)] to volatiles of peach wood, either natural or chemically fractionated, were observed. Mated females were stimulated by and responsive to such materials and deposited significantly more eggs on substrates, including unnatural hosts, that had been treated with aqueous mixtures of bark-canker materials. Stim ulation to oviposit occurred even when the female was blinded, indicating the presence of chemical cues. Natural canker-bark extracts immediately stimulated ovipostion and for a few hours significantly increased the number of eggs laid. However, average fecundity was not increased. Antennectomy did not significantly decrease response to volatiles by gravid females, and alternate sites of such chemoreception were not located. Complex mixtures derived by solvent extraction, steam distillation, and volatiles trapping from bark, canker, and gum all had activity. Observations of insect behavior in outdoor cages and also in the laboratory indicated that visual, chemosensory, and mechanosensory receptors are involved in host finding and oviposition. Key Words-Attractants, extracted volatile compounds, Synanthedon pic tipes, Lepidoptera, Sesiidae, chemosensory, insect behavior, oviposition, mechanosensory, host-insect interactions, olfactory receptors, oviposition behavior, apple, peach, pear, plant odors, Prunus spp. I Mention of firm or product names does not imply recommendation or endorsement by USDA over others not mentioned. 2 Present address: Asian Parasite Laboratory, USDA, ARS, c/o American Embassy, Seoul, Korea, APO San Francisco 9630I. 237 0098-0331/88/0100-0237 06.00/0 © 1988 Plenum Publishing Corporallon 238 REED ET AL. I TRODUCTIO In recent years, research on elucidation of insect-host interactions has included ovipositional responses of gravid females to their hosts. This relationship is crucial in most insect-plant interactions, but unfortunately it is sometimes dif ficult to establish and is the least understood relationship involved in such in teractions (Stadler, 1983). It is generally accepted that chemical cues may in voke both long- and short-range orientation by the ovipositing female to the host. The distinction between feeding and ovipositional stimulation is often ob scure, as exemplified by the dipteran pests (Liriomyea spp.). Fortunately, in studying nonfeeding lepidopterous females this problem is not encountered. However, with such females, moisture may become extremely important in their host-seeking (Saxena and Goyal, 1978). Another issue that needs clarifi cation is the distinction between ovipositional stimulation and ovipositional at traction. Although these functions are not necessarily mutually exclusive (Brown et al., 1970), a chemical attractant may not stimulate oviposition, and, con versely, a stimulant may not attract. The lesser peachtree borer, Synanthedon pictipes (G&R) (LPTB), is an important pest of peaches and cherries in most production areas. The host range of LPTB consists of the genus Prunus; it may therefore be considered a re stricted feeder, thus requiring fairly specific host-finding mechanisms. Females tend to oviposit on roughened bark of healthy trees near wounds where fresh gum is present (Bobb, 1959). Neonate larvae utilize light, chemotaxes, and thigmotaxes as cues in an active search for entry into the bark (Wiener and Norris, 1982). The reproductive behavior of adult Sesiidae females has been reported by Girault (1907), King (1917), Bobb (1959), and Cleveland et al. (1968), but the chemical relationships between the mated female and her host plants have not been defined. Observations have generally concluded that fe males are markedly attracted to the gummy deposits on peach wood that orig inate from cankers formed by cold damage, Cytospora sp. fungi, or mechanical injuries. A chemical isolated from peach bark, larval frass, and gum mixtures was reported by Gentry and Wells (1982) to be attractive to gravid females of peachtree borer, S. exitiosa (Say). Since S. exitiosa and S. pictipes are conge neric species, some commonality in such behavior might be expected. Our research was conducted to quantify the attractancy of gravid LPTB females to components of peach bark and to determine if such components stim ulate oviposition or increase fecundity. This paper presents the results of our attempts to isolate and identify such components and to record the ovipositional behavior of LPTB in the laboratory and field at Vincennes, Indiana, Peoria, Illinois, and Delaware, Ohio. OVIPOSITIO AL BEHAVIOR OF PEACHTREE BORER 239 METHODS AD MATERIALS LPTB was reared using the methods of Cleveland et al. (1968). Newly mated females were maintained in screen cages on damp sand at 10 DC until used in experiments, either the same or the next day. Various parameters of response were determined in preliminary experiments. For experiments utiliz ing natural canker and gum, the cankerous areas on either Cresthaven or Red skin peach trees were removed by cutting both sides and ends of the canker and scraping bark, canker, gum and debris from the entire area. Gum was isolated by carefully selecting debris-free material. Healthy bark used as check material was peeled from 10- to 15-cm-diam. limbs from the same trees. All test mate rials were maintained in sealed plastic bags at 4 ± 1DC until used. Materials for scanning electron microscopy were mounted on aluminum stubs with con ductive cement and sputter-coated (model Hummer V, Technics, Springfield, Virginia) with 500 A of gold. SEM observations were performed with a Hitachi model S-500 (Mountainview, California) at 20 KV accelerating voltage. Chemical Cues Experiment. Small branches (2.5 x 25 cm), cut from pear trees (a LPTB nonhost) and from peach trees (a host) were presented to LPTB adult females in 50 x 50 x 50-cm screen cages in an air-conditioned green house. In test 1, branch ends were impaled by nails to a 35 x 35-cm board and exposed to 10 gravid females. Pear branches were coated with aqueous canker gum mixtures and tested in comparison to nontreated (clean) pear branches and water wicks. In test 2, peach wood was similarily treated and exposed to LPTB. The board was rotated periodically to ensure equal environmental exposure of each treatment. The test was replicated six times and data were submitted to analysis of variance. Visual Cues Experiments. In test 1, to determine the role that sight may play in host response, latex paint was applied to both compound eyes of 10 mated females. No adverse effects of the paint treatment upon the insects was noted. These treated insects were placed into a 30 x 30 x 30-cm screen cage which contained one peach branch (10 x 2 cm), treated with an aqueous canker gum-bark mixture from Redskin peach trees, and one pear branch the same size. A duplicate cage was used with 10 control (nonpainted) females. Behavior and egg deposition were observed in both cages for 6 hr. In Test 2, females, either normal or blinded with latex paint, were caged with apple, pear, and peach branches (10 x 2 cm). The canker-gum-bark mix ture was layered onto a moist cotton pad placed beneath the branches but in accessible to the insects. The cages consisted of 3. 78-liter ice-cream cartons (20 x 18 cm) with the middle of the carton removed and replaced with screen wire. A screen bottom separated the cotton pad 5 cm from the rest of the cage. 240 REED ET AL. Treated cages as well as controls without the canker material consisted of three replications of three insects. After log transformation, 24-hr egg counts were submitted to analysis of variance and Duncan's (1955) multiple-range test. Ovipositional Stimulation Experiments. Two other types of tests were con ducted to determine if peachwood canker or gum might actively stimulate ovi position or increase fecundity. In the first experiment, air from the exhaust of an air compressor was passed through flasks containing aqueous suspensions of either canker material, gum, healthy bark, or distilled water. The volatile-laden air immediately flowed into four 13.5 x 10.5 x 3.5-cm plastic sweater boxes containing exhaust ports in the opposite ends. The boxes were layered with 2 cm-thick cotton batting saturated with distilled water. Twelve females, eclosed and mated the same day, were introduced into each box, and the eggs that were deposited on either the cotton or the box were counted after 24 hr in one series of tests and 48 hr in a second series of tests. Another experiment was conducted in individual plastic IOO-ml cups (6.5 x 6 cm) with copper screen covers. Aqueous canker material (0.5 ml) was applied to the center of a I-cm layer of moist cotton either on top of the cotton so that the insects could contact it or inverted so that the material was not accessible either tactilely or visually to the insects although they were exposed to volatiles. Each treatment was replicated 25 times; moist cotton pads were used as controls. Surgically altered LPTB females were used as subtreatments consisting of five replicate Latin-square arrangements as follows: (1) normal mated female, (2) mated female with one antenna removed at base of pedicel, (3) mated female with both antennae removed, (4) mated female with both an tennae three-fourths removed, and (5) mated female with tip of hypopharnyx removed. Cups were maintained in an air-conditioned greenhouse at 85 ± 5°C and 75 ± 10% relative humidity. Eggs were counted at 24-hr intervals for 72 hr.