Age-related responses from the maxillary sensilla styloconica of Choristoneura fumiferana larvae to foliage extracts from balsam fir hosts

An electrophysiological study of the sensilla styloconica of the galea in Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae) larvae showed a differential response between fourth- and sixth-instars to extracts of balsam fir foliage. Larvae raised on artificial diet were stimulated with the water soluble fraction of needle extracts obtained from terminal and lateral shoots of 30- and 70-yr-old balsam fir trees. An extract-sensitive neuron was found in the lateral styloconic sensillum of both instars. The lateral styloconica in the fourth-instar larvae were more sensitive to extracts from terminal than from lateral shoot foliage of both young and old trees. The lateral styloconica of sixth-instar larvae were more sensitive to lateral shoot foliage of old trees. Results are discussed with respect to their relationship to feeding preferences and feeding rates observed in a previous behavioural study.     

Different sensitivities of the sugar receptor of the lateral styloconic sensillum in fourth- and sixth-instar larvae of the spruce budworm Choristoneura fumiferana

Female fourth- and sixth-instar larvae, Choristoneura fumiferana, were tested individually for the response of the sugar cell on the lateral styloconic sensillum to 25 mM/l concentrations of 12 carbohydrates. The spruce budworm showed an age-related change in responsiveness of the sugar cell. The order of stimulating effectiveness for fourth-instars was melibiose > sucrose > raffinose.These storage di- and trisaccharides are present in the host plant at the beginning of budbreak. Sixth-instars responded to sucrose > fructose> m-inositol. These findings are in accordance with those of a previous behavioural study on feeding preferences of sixth-instars. The response for both melibiose and raffinose does not change from fourth- to sixth-instars; however, it does for sucrose, fructose and m- inositol.     

Detection of alkaloids and carbohydrates by taste receptor cells of the galea of gypsy moth larvae, Lymantria dispar (L.)

Gypsy moth larvae are polyphagous feeders. The electrophysiological responses of the medial and lateral styloconic sensilla to four secondary compounds (e.g., alkaloids), two carbohydrates, and one inorganic salt were examined using an extracellular tip-recording method. In the medial sensillum, one taste receptor cell responded to the alkaloids, strychnine, caffeine, nicotine, and aristolochic acid (i.e., deterrent-sensitive cell), while another, responded to the sugar alcohol and inositol (inositol-sensitive cell). In both medial and lateral sensilla, two taste receptor cells in each sensillum responded minimally and sporadically to 30?mM potassium chloride (KCl) (i.e., KCl-sensitive cells); one cell produced much larger amplitude action potentials than the other. In the medial sensillum, only the large-amplitude KCl-sensitive cell exhibited an increased firing rate with increasing salt concentration. When binary mixture experiments were conducted, it was confirmed that the large-amplitude KCl-sensitive cell and the deterrent-sensitive cell in the medial sensillum were one in the same cell. Only a single cell in the lateral sensillum responded to the sugar, sucrose (sucrose-sensitive cell). The temporal dynamics of responses of the deterrent-sensitive, sucrose-sensitive, and inositol-sensitive cells were compared. Concentration?Cresponse data were obtained for the deterrent-sensitive cell to various alkaloids, as well as to KCl.     

Taste cell responses in the polyphagous arctiid, Grammia geneura: towards a general pattern for caterpillars

The caterpillars of Grammia geneura are polyphagous as individuals. Electrophysiological responses of its medial and lateral galeal styloconic sensilla to 21 amino acids, 6 carbohydrates, 10 chemically diverse plant secondary compounds and two inorganic salts were examined. In the medial sensillum, a single cell responded to 8 amino acids, 3 carbohydrates, and the iridoid, catalpol, which is present in a favored hostplant. In the lateral sensillum, one cell responded to amino acids and another to fructose. Two cells in each sensillum responded to secondary compounds and it is suggested that the same cells are stimulated by inorganic salts. There was no evidence of a separate salt-sensitive cell. Phenylalanine stimulated a deterrent cell in the medial sensillum and was behaviorally deterrent. Some essential amino acids did not stimulate any cells and it is suggested that a small number of amino acids (sometimes non-essential) may serve as indicators of nutrient quality. Sugars probably serve as the primary phagostimulants because they are in relatively high concentrations in plants. It is proposed that taste receptor cells should be categorized primarily by their behavioral effects as phagostimulatory or deterrent, rather than their specific ranges of responsiveness. This would emphasize the basic similarities across taxa.     

Responses of the African Armyworm and three species of borers to carbohydrates and phenolic substances: an electro- and behavioural physiological study

Electrophysiological recordings were made from the maxillary sensilla styloconica of fifth instars of the African Armyworm, Spodoptera exempta (Wlk.) (Lep.: Noctuidae), and the borers (Lep.: Pyralidae) Eldana saccharina Wlk. and Maruca testulalis (Geyer), and fifth and sixth instars of the borer Chilo partellus (Swinhoe). Stimuli were different concentrations of 13 carbohydrates and 11 phenolic substances. It appeared that a sucrose sensitive cell is present in the lateral styloconic sensilla of Spodoptera, Maruca and Chilo, and in both the lateral and medial sensilla of Eldana. The sucrose cell of fifth-instar Chilo is more sensitive than that of sixth instars. The additional responses of the medial cells to the phagostimulant sucrose are supposed to have contributed to the widespread infestation of sugarcane by Eldana. The medial sensilla of the Armyworm contain a cell type responding to both meso-inositol and D-ribose. Chilo appeared to be the only one of the three borer species able to perceive phenolic substances. It has a cell responsive to chlorogenic acid in its medial sensilla, which is a bit more sensitive in fifth- than in sixth-instar larvae. The presence of this cell may account for the positive correlation between total phenols content and resistance of sorghum cultivars to this borer. Spodoptera can perceive four phenolic acids. Its lateral sensilla contain a cell responsive to chlorogenic acid and its medial sensilla a cell responsive to 4-hydroxybenzoic acid, gallic acid as well as vanillic acid. Feeding experiments showed that the latter substances inhibit feeding in the Armyworm.So far, in the borers adequate stimuli could be determined for only tow or three of the four taste cells present in each sensillum styloconicum. This indicates that we are still unaware of chemicals that may be important to these caterpillars. On the other hand, for the sensilla styloconica of Spodoptera, electrophysiology suggests the presence of five or six taste cell types, whereas only four taste cells are known from morphological studies.     

Fine structure of the galeal styloconic sensilla of larval Lymantria dispar (Lepidoptera: Lymantriidae)

Lepidopteran larvae possess two pairs of styloconic sensilla located on the maxillary galea. These sensilla, namely the lateral and medial styloconic sensilla, are each comprised of a smaller cone, which is inserted into a style. They are thought to play an important role in host-plant selection and are the main organs involved in feeding. Ultrastructural examination of these sensilla of fifth instar Lymantria dispar (L.) larvae reveal that they are each approximately 70 um in length and 30 um in width. Each sensillum consists of a single sensory peg inserted into the socket of a large style. Each peg bears a slightly subapical terminal pore averaging 317 nm in lateral and 179 nm in medial sensilla. Each sensillum houses five bipolar neurons. The proximal dendritic segment of each neuron gives rise to an unbranched distal dendritic segment. Four of these dendrites terminate near the tip of the sensillum below the pore and bear ultrastructural features consistent with contact chemosensilla. The fifth distal dendrite terminates near the base of the peg and bears ultrastructural features consistent with mechanosensilla. Thus, these sensilla each bear a bimodal chemo-mechanosensory function. The distal dendrites lie within the dendritic channel and are enclosed by a dendritic sheath. The intermediate and outer sheath cells enclose a large sensillar sinus, whereas the smaller ciliary sinus is enclosed by the inner cell. The neurons are ensheathed successively by the inner, intermediate, and outer sheath cells.     

草地贪夜蛾幼虫对四种刺激物质的味觉感受和取食选择

【目的】为了筛选有效的草地贪夜蛾Spodoptera frugiperda幼虫取食激食素和抑制剂并探究其味觉感受机理,为生态防治草地贪夜蛾提供理论和实践上的依据。【方法】利用单感受器记录法测定草地贪夜蛾5龄第2天幼虫下颚外颚叶上中栓锥感器和侧栓锥感器对不同浓度的蔗糖、黑芥子苷、单宁酸和盐酸奎宁4种刺激物质的电生理反应,并采用二项叶碟法测定草地贪夜蛾幼虫对这些刺激物质的取食选择行为。【结果】草地贪夜蛾幼虫中栓锥感器和侧栓锥感器内均存在对蔗糖、黑芥子苷和单宁酸敏感的味觉受体神经元,但是神经元的活性随着刺激物的种类及浓度而变化。其中,两类感器内神经元对蔗糖和黑芥子苷的反应均呈现典型的浓度梯度反应。中栓锥感器内存在对盐酸奎宁敏感的味觉受体神经元,但是呈现逆浓度梯度的反应模式,侧栓锥感器内不存在对盐酸奎宁敏感的神经元。蔗糖显著诱导幼虫的取食行为,而盐酸奎宁、黑芥子苷和单宁酸均抑制幼虫的取食行为,且都呈现浓度梯度的抑制活性。【结论】草地贪夜蛾幼虫中栓锥感器和侧栓锥感器内均存在对取食激食素和抑制剂敏感的味觉受体神经元,但是两类感器不论在反应谱上还是敏感性上均存在差异。蔗糖可以作为取食激食素,盐酸奎宁...     

Electrophysiological responses to sucrose from a gustatory sensillum on the larval maxillary palp of the spruce budworm,Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

The tip of the maxillary palp in the spruce budworm and other lepidoptera larvae contains a cluster of eight basiconic sensilla. Except for a few electrophysiological recordings from the entire group of these sensilla, no information is available on the response characteristics of any individual one. Using a compound microscope at 320x magnification, with a long working-distance objective, it has been possible to view individual sensilla and to record electrical responses from three of these to some gustatory stimuli in preliminary tests. One of them, sensillum (L1), contains a sugar-sensitive neuron whose responses to a range of sucrose concentrations are reported here. The physiological characteristics of this neuron differ from those of the known sugar-sensitive neuron in the lateral styloconic sensillum on the galea of these same insects. Input of chemosensory information about the same gustatory stimulus by more than one neuron in different mouthpart sensilla is discussed in relation to the feeding habits of the spruce budworm.     

Food selection in adults and larvae of three species of Lepidoptera: a behavioural and electrophysiological study

Adults and larvae of Spodoptera littoralis, Heliothis virescens and Heliothis armigera were tested with a range of sugars, amino acids, sugar alcohols and allelochemics. Feeding behaviour was correlated with the electrophysiological responses of maxillary styloconic sensilla in the larvae and proboscis styloconic sensilla in the adult. The neural response was more vigorous in larvae than in adults but otherwise the spectra of responses were similar in the two life stages. Phagostimulants and antifeedants stimulated maximally different sensilla in larvae but not in adults. The responses of adult sensilla to sugars and amino acids were significantly correlated to behaviour in all three species, but only in H. armigera was there a similar correlation with the sugar alcohols and allelochemics.     

Chemoreception of amino acids in larvae of two species of Pieris

ABSTRACT. Specificity and sensitivity of gustatory neurones in response to twenty-two amino acids were studied in larvae of Pieris brassicae L. and Pieris rapae L. (Lepidoptera: Pieridae) using electrophysiological methods. Twelve amino acids stimulated a specific amino acid receptor cell in the lateral styloconic sensillum on the maxillary galea of both species, and a further two evoked single unit responses in the same sensillum of P.brassicae only. Histidine, phenylalanine and tryptophane were the weakest stimulants for P.brassicae , but were among the four best stimulants for P.rapae . In both species, eight amino acids were ineffective. Significant differences in stimulatory effectiveness were found between amino acids. Nutritionally essential amino acids were more effective in both species, as in five other lepidopterous species. Similarities with postulated sites for amino acid recognition in the dipteran Boettcherisca peregrina were found.
Concentration-response (C/R) relations were studied for five amino acids. Significant differences were found in saturated response levels. Parameters characterizing C/R relations were estimated using a logistic model. Comparing C/R parameters with phytochemical data on concentrations of free amino acids in a common host plant, Brassica oleracea L., shows that amino acids are effective stimuli at their natural concentrations. The amino acid chemoreceptor seems able to transmit information about concentration differences of amino acids in the plant tissue.     

Electrophysiological responses of taste cells to nutrient mixtures in the polyphagous caterpillar of Grammia geneura

In a normally feeding insect, the taste receptors are exposed to complex mixtures of chemicals, not single compounds. We investigate the responses of neurons in the galeal sensilla of the caterpillar of Grammia geneura to mixtures of nutrient compounds at concentrations occurring in plants. Compounds that stimulated the same neuron were generally additive in their effects in binary mixtures. Amino acids that did not stimulate usually had no effect in mixtures with a stimulating compound, but glutamic acid reduced the response to serine in the medial sensillum. Nutrient compounds that stimulated different cells in a sensillum acted independently of each other. Complex mixtures of amino acids resembling samples of free amino acids from three host plants were less stimulating than expected from their molar concentrations. In host plant selection, the response from the medial sensillum is probably dominated by sucrose; unless sucrose levels are low, amino acids will contribute little to sensory input because they stimulate the same cell as sucrose. In the lateral sensillum, amino acids act independently of sugars. The limited contact chemosensory array of caterpillars seems inadequate to allow them to make fine distinctions between plants on the basis of their free amino acids.     

Distribution of chemo- and mechanoreceptors on the antennae and maxillae of Busseola fusca larvae

The stem borer Busseola fusca (Fuller) (Lepidoptera: Noctuidae) is a major pest of maize, Zea mays L., and sorghum, Sorghum bicolor (L.) Moench (both Poaceae), in sub-Saharan Africa. Like in many other lepidopteran insects, the success of B. fusca in recognizing and colonizing a limited variety of plants is based on the interaction between its sensory systems and the physicochemical characteristics of its immediate environment. The sensilla on the maxillary galeae of B. fusca larvae are typical of Lepidoptera and comprise two uniporous styloconic sensilla, which are contact chemoreceptors, three basiconic sensilla, and two aporous sensilla chaetica. The maxillary palp is two-segmented and has eight small basiconic sensilla at the tip, which were also found to be gustatory. The antennae of B. fusca larvae are short and simple. The sensilla of the antenna are composed of two aporous sensilla chaetica, three multiporous cone-shaped basiconic sensilla, three small basiconic sensilla, and one aporous styloconic sensillum. The basiconic sensillum located on the third antennal segment displayed a contact chemoreception response. The other basiconic sensilla did not show any action potential activity in tip-recording tests. The significant and positive dose–response curve obtained for the antennal basiconic sensillum with sucrose indicated for the first time the presence of gustatory chemoreceptors on the antennae of a lepidopteran larva.     

Gustatory perception of phytoecdysteroids in Plodia interpunctella larvae

Phytoecdysteroids are steroidal compounds produced by various plants that disrupt growth and development of insects eating them. They exhibit an insecticidal activity on a number of insect pests such as Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae). In this study, we further evaluated whether phytoecdysteroids deter larvae of this species from feeding, by using four phytoecdysteroid molecules, commonly occurring in plants: 20‐hydroxyecdysone (20E), ponasterone A (PonA), polypodine B (PolB), and makisterone A (MakA). Fourth instar P. interpunctella avoided contact with food pellets treated with these phytoecdysteroids in a dose‐dependent way (2–30 mm ). In order to test whether this avoidance was mediated by taste sensitivity, we recorded the responses of taste neurons located in the lateral and medial sensilla styloconica of the galea. At least one neuron responded to each of these compounds in both sensilla. The neuron located in the medial sensillum had a detection threshold of 10^?6 m for PonA, 10^?4 m for 20E and PolB, and 10^?3 m for MakA. The lateral sensillum neuron responded with less intensity and its detection threshold was 10 times higher than that of the medial sensillum neuron. These results indicate that phytoecdysteroids are detected as deterrent stimuli by P. interpunctella larvae and that small structural differences significantly affect their biological activity.     

Flavonoids from cabbage are feeding stimulants for diamondback moth larvae additional to glucosinolates: Chemoreception and behaviour

In caterpillars two styloconic contact chemoreceptors on the maxillary galea are assumed to contain the main taste receptors involved in host plant selection. The diamondback moth, Plutella xylostella L. is a specialist feeder of plants in the Brassicaceae, a plant family characterized by the biosynthesis of glucosinolates. We used pea (Pisum sativum L., Leguminosae) as a neutral non-host for a dual-choice leaf disc assay to quantify feeding stimulation by glucosinolates and flavonoids. Increasing concentrations of sinigrin resulted in significant preferences for sinigrin-treated leaf discs, with a threshold between 1 and 3 M. Millimolar concentrations of four of the five flavonol triglucosides likewise elicited a significant preference for flavonoid-treated leaf discs. A mixture of four flavonoids and sinigrin was significantly preferred over sinigrin-treated leaf discs alone. Vigorous unicellular electrophysiological responses of medial maxillary styloconic taste sensilla were observed in response to five glucosinolates (glucocapparin, sinigrin, glucobrassicin, glucoiberin, and gluconasturtiin). This medial taste neuron responded in a dose-dependent manner to a concentration series of sinigrin, with a threshold of response of ca. 1 M. The lateral sensillum styloconicum contained a neuron sensitive to sucrose, glucose, and fructose. However, no responses in the two types of maxillary styloconic sensilla to the phagostimulatory flavonoids could be detected, suggesting that other taste organs mediate chemoreception of flavonoids. We conclude that diamondback moth larvae employ a combination of biosynthetically distinct categories of feeding stimulants which allows for a higher degree of discriminatory ability than when this would be based on glucosinolates alone.     

Analysis of the electrical responses of antennal thermo- and hygroreceptors of Antheraea (Saturniidae,Lepidoptera) to thermal,mechanical, and electrical stimuli

Summary Antennal styloconic thermo-hygro sensilla of Antheraea were studied with DC-coupled transepithelial recordings. — The transepithelial voltage changed by about 2 mV · °C^–1. The spike frequency of the cold cell reached 300 Hz at the onset of negative temperature steps, but only 30 Hz at static temperatures (as with metal electrodes). The cold cell spikes showed a brief afterhyperpolarization that increased with temperature. The spikes of the cold- and warm-stimulated cells facilitated each other at low temperature. Mechanical stimuli (push against the sensillum, hydrostatic pressure of < ± 50 kPa, ultrasonic vibrations 120 kHz) modified the responses of the cold- and the warm-stimulated cells. Latency of cold cell responses to ultrasonic stimulation was occasionally less than 3 ms. — The impulse frequencies of the warm and the cold cells depend on the temperature and the magnitude of temperature change. When the firing rate is high enough by either or both of these parameters, it can be forced still higher by application of clamp current (outside positive). The higher the firing rate prior to clamping, the greater the effect of the current. — By analogy with sensilla for other modalities, this relationship between frequency and clamp current strongly suggests that stimulus-dependent changes in the conductance of dendritic membranes control the excitation of the warm and cold cells.Abbreviations DC direct voltage - TER transepithelial slope resistance between recording electrode and reference electrode in the hemolymph - NTC thermistor with negative temperature coefficient - TEV transepithelial voltage between electrodes - THS thermo-hygro sensillum     

Comparative external ultrastructure and diffusion pathways in styloconic sensilla on the maxillary galea of larval Mamestra configurata (Walker) (Lepidoptera: Noctuidae) and five other species

The external ultrastructure of sensilla on the maxillary galea are investigated in Mamestra configurata and five other lepidopterous larvae using scanning electron microscopy. The galea and lacinia, comprising the mesal lobe of the maxilla, are either completely separate, fused, or incompletely fused in these species. The distal surface of the mesal lobe of all species examined bears two styloconic sensilla, three basiconic sensilla, and three trichoid sensilla, whereas the midventral wall of this lobe bears a campaniform sensillum. The latter sensillum is visible in only three of the six species examined. The styloconic and basiconic sensilla occupy a ventro anterior location, whereas the trichoid sensilla are positioned dorsoposteriorly. Interspecific comparisons of galeal size, as well as sensillar size, shape, and position are made for all species. The styloconic sensilla are the only sensillar type permeable to an aqueous solution of cobalt chloride when viewed by brightfield light microscopy in all species examined. Cobalt ions permeate through the terminal pore of each styloconic peg and percolate through the fenestrated fibrillar pore matrix, located directly below the pore. These ions permeate along the dendritic channel and accumulate in the adjacent sensillar sinus surrounding the peg and/or style by way of a presumably permeable dendritic sheath in all species, but to varying extents. The cuticular sidewall pores surrounding the terminal pore also appear to be permeable to cobalt ions in all the species examined. In most species examined, the styloconic sensilla are only minimally permeable to mercury ions. In these species, mercury ions permeate through the terminal pore, but become trapped within the plug of fenestrated fibrils within it. The sidewall pores are not permeable to mercury ions in any of the species examined. The styloconic sensilla are not permeable to lead ions in M. configurata or Malacosoma lutescens, the only species tested. © 1996 Wiley-Liss, Inc.     

Physiology of a sucrose-sensitive cell on the galea of the eastern spruce budworm larva, Choristoneura fumiferana

ABSTRACT The lateral styloconic sensillum on the galea of the eastern spruce budworm larva Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae) contains a cell which responds to sucrose. The electro-physiologial threshold for response is below 0.5 mM sucrose. The K_b for the response is 1.5 mM, and the V_max is 200 impulses/s. The physiological data are interpreted with respect to the sucrose-stimulated feeding behaviour of the larva.     

Multiple mode of action of the feeding deterrent,toosendanin, on the sense of taste in Pieris brassicae larvae

Toosendanin, a tetranortriterpenoid isolated from the bark of Melia toosendan, is a feeding deterrent for larvae of Pieris brassicae. By using electrophysiological techniques, it was found that toosendanin stimulates a deterrent receptor cell located in the medial maxillary sensillum styloconicum. Toosendanin also inhibits responses of both the sugar and glucosinolate receptor cell, which are localized in the lateral sensillum styloconicum. The degree of inhibition of the sugar receptor increases with increasing sucrose concentration. The glucosinolate receptor cell shows a reversed reaction: inhibition by toosendanin decreases with increasing sinigrin concentration. Inhibitory effects occur at a toosendanin concentration as low as 10^–9 M and are dose dependent. The taste neurons that respond to amino acids or deterrents in the lateral sensillum, however, are not affected by toosendanin. It is concluded that the sensory code underlying feeding behaviour is modulated by toosendanin via several different peripheral sensory mechanisms.     

Physiological differences between two sugar-sensitive neurons in the galea and the maxillary palp of the spruce budworm larva Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

The L1 contact-chemoreceptor sensillum on the maxillary palp of the spruce budworm larva Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) was examined electrophysiologically for its responses to stimulation by various pyranose and furanose sugars. The results were compared to those from previous work on the sugar-sensitive neuron of the LST sensillum on the galea. We show that the L1 contact-chemoreceptor sensillum contains one sugar-sensitive neuron with furanose but no pyranose sites. It has response characteristics that differ from those of the sugar-sensitive neuron in the LST. Behavioural 2-choice feeding experiments show that, even with both known sugar-sensitive neurons disabled, larvae can still discriminate between disks treated with either distilled water or alpha-D-glucose. We conclude that the epipharyngeal sensilla must thus also contain a sugar-sensitive neuron.     

An electrophysiological analysis of the effect of phagostimulant mixtures on the responses of a deterrent-sensitive cell of gypsy moth larvae, Lymantria dispar (L.)

Gypsy moth larvae, Lymantria dispar (L.), are polyphagous feeders. The medial styloconic sensillum of this species bears a taste receptor cell that responds to alkaloids and another that responds to the sugar alcohol, inositol. The lateral styloconic sensillum bears a taste receptor cell that is sensitive to the sugar, sucrose. We tested the effect of two phagostimulants, namely sucrose and inositol, on the response of the deterrent-sensitive cell and found that both phagostimulants suppressed its response, equally, while their combination was significantly more effective. We also tested the effect of two alkaloids (i.e., strychnine and caffeine), which deter feeding in this species, on the response of the inositol- and sucrose-sensitive cells. Although both of these deterrents had no effect in suppressing the response of the sucrose-sensitive cell, they both had an effect in suppressing the inositol-sensitive cell. We also found that sucrose suppressed the response of the inositol-sensitive cell, whereas inositol had no significant effect on the response of the sucrose-sensitive cell. In this paper, we examined the effect of mixtures of these compounds to determine the nature of their interaction. In the context of host–plant interactions and, for example, host recognition, whereby host plant acceptability depends on the total sensory impression acquired from responses to multiple plant components rather than the presence or absence of single stimulant or deterrent compounds, this study could have a direct bearing in the development of natural compounds (i.e., alkaloids) for pest control and crop protection. It will also contribute to our understanding of the neural basis of the feeding behavior of this insect.