Electrophysiological characterization of olfactory cell types in the antennae and palps of the housefly

A set of odours was presented to the housefly Musca domestica and the electrophysiological responses of single olfactory receptor cells in the antennae and palps were recorded. The olfactory cells in the antennae of the housefly showed a large variability of response profiles, but multidimensional cluster analysis suggested a moderate clustering in olfactory response types. Receptor cells with similar or with different odour response profiles can reside in one and the same sensillum. No fixed spatial distribution of olfactory response types over the antennal of palpal surface was found. The odours of 1-octen-3-ol, amyl acetate, 3-methylphenol, 2-pentanone and R(+)limonene elicited the largest responses in antennal cells. Most odours elicited responses in cells of only a few of the clusters, but 1-octen-3-ol was detected by cells of almost all clusters of the antenna. Surprisingly, rather low responses were found to acetic acid, skatole, indole and muscalure, odours that are known to attract flies. Response profiles of palpal cells differed considerably from those of antennal cells. Palpal cells mostly responded to 3-methylphenol and 2-pentanone. In the palps, the clusters of cells responding to 3-methylphenol and 2-pentanone are clearly separated from the other olfactory cells.     

Morphological and functional heterogeneity in olfactory perception between antennae and maxillary palps in the pumpkin fruit fly,Bactrocera depressa

The morphology and ultrastructure of the olfactory sensilla on the antennae and maxillary palps were investigated through scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and their responses to five volatile compounds were measured using electroantenogram (EAG) and electropalpogram (EPG) techniques in the pumpkin fruit fly, Bactrocera depressa (Shiraki; Diptera: Tephritidae). Male and female B. depressa displayed distinct morphological types of olfactory sensilla in the antennae and maxillary palps, with predominant populations of trichoid, basiconic, and coeloconic sensilla. Basiconic sensilla, the most abundant type of olfactory sensilla in the antennae, could be further classified into two different types. In contrast, the maxillary palps exhibited predominant populations of a single type of curved basiconic sensilla. High‐resolution SEM observation revealed the presence of multiple nanoscale wall‐pores on the cuticular surface of trichoid and basiconic sensilla, indicating that their primary function is olfactory. In contrast, coeloconic sensilla displayed several longitudinal grooves around the sensillum peg. The TEM observation of individual antennal olfactory sensilla indicates that the basiconic sensilla are thin‐walled, while the trichoid sensilla are thick‐walled. The profile of EAG responses of male B. depressa was different from their EPG response profile, indicating that the olfactory function of maxillary palps is different from that of antennae in this species. The structural and functional variation in the olfactory sensilla between antennae and maxillary palps suggests that each plays an independent role in the perception of olfactory signals in B. depressa.     

Olfactory and behavioural responses of tabanid horseflies to octenol,phenols and aged horse urine

Electrophysiological and behavioural responses of females of two tabanid species, Tabanus bromius L. and Atylotus quadrifarius (Loew) (Diptera: Tabanidae), to ammonia, octenol (1‐octen‐3‐ol), phenols and aged horse urine were compared. Electroantennogram (EAG) responses in both species to octenol, 4‐methylphenol (4MP), 3‐propylphenol (3PP) and a phenol mixture (4MP and 3PP at a ratio of 16 : 1) increased in a dose‐dependent fashion. The most effective stimulus was 4MP and synergism between the two phenols may exist. Aged horse urine also elicited strong EAG responses in both species. Using gas chromatography–mass spectrometry (GC‐MS) analysis, we identified 29 compounds in horse urine, which included, in particular, ketones, fatty alcohols and phenols, among which 4MP was the most abundant component (? 80%). Trapping experiments were carried out using Nzi traps baited with various odours. Octenol and the phenol mixture in combination with ammonia increased catches of tabanids by 1.8–2.8 times relative to ammonia alone. Aged horse urine increased catches of T. bromius and A. quadrifarius by 2.2 and 4.1 times, respectively. The high attractiveness of aged horse urine, especially for A. quadrifarius, is not likely to derive from 4MP alone, but from the mixture of various active compounds used in host location.     

Quantifying relative importance of maxillary palp information on the olfactory behavior of Drosophila melanogaster

Maxillary palps have been proposed as secondary olfactory organs, after the antennae, in Drosophila melanogaster. Our study tries to establish the quantitative importance of both organs as olfactory information mediators. Dose-response curves for three odorants: ethyl acetate, propionaldehyde and benzaldehyde were carried out for comparing olfaction in either complete animals or flies surgically deprived of antennae. Antennaless flies tested in our behavioral assay showed indifferent, attractant and repellent responses depending on concentration, similarly as normal flies do. However, they clearly displayed less sensitivity than normal flies. The range of concentrations they were able to perceive was correlated to antennal sensitivity approximately by a factor 110 for ethyl acetate and benzaldehyde, and between 110 and 1100 at high concentrations of propionaldehyde. A complementary experiment was performed to test changes in olfactory behavior produced by removing maxillary palps in the presence of antennae. At high concentrations of odorant, responses to ethyl acetate and propionaldehyde experienced small changes when both palps were removed. Results are compatible with a summation model of all olfactory information reaching the brain either through antennae or palps.Abbreviations ANOVA analysis of variance - EAG electroantennogram, extracellular recording of electrical changes produced on the antenna in response to odorant stimulation - EPG electropalpogram, extracellular recording of electrical changes produced on the maxillary palp in response to odorant stimulation - IO Olfactory index     

The role of the antennae and maxillary palps in mediating food preference by larvae of the tobacco hornworm, Manduca sexta

The role of antennae and maxillary palps in mediating food preferences by Manduca sexta (Johan.) (Lepidoptera: Sphingidae) was examined. Fifth‐instar larvae, having different chemosensory organs remaining after microsurgery, were tested in two‐choice feeding assays using three solanaceous host species Lycopersicon esculentum (Mill.), Datura innoxia (Mill.), and Solanum pseudocapsicum (L.), an acceptable non‐host Raphanus sativus (L.) (Cruciferae), and an unacceptable non‐host Pelargonium hortorum (L.H. Bailey) (Geraniaceae). Larvae had a choice between leaf discs of two plant species or between one species and moist filter paper discs (water). The antennae are fully competent in mediating normal (unimpaired) food preferences for S. pseudocapsicum vs. water and P. hortorum vs. water. Thus, the antennae alone can mediate both acceptance and rejection behaviour. The latter is the first report of such a function in M. sexta. The antennae are partly competent (reduced preference) for S. pseudocapsicum vs. P. hortorum. No antennal competence could be demonstrated using the other plant species tested. The antennae alone are either partly needed (R. sativus vs. water) or not necessary to elicit normal food preferences for the plant species tested. The maxillary palps are fully competent in mediating normal food preference for S. pseudocapsicum vs. water, and D. innoxia vs. water. The palps are partly competent for S. pseudocapsicum vs. P. hortorum and incompetent for the other plant species tested. Thus, the maxillary palps alone could only mediate acceptance behaviour in this study. The need of the maxillary palps in normal food preferences could not be demonstrated. Interestingly, the maxillary palps alone can mediate food preferences for two normally rejected plants, R. sativus and P. hortorum. Similar results are reported for two other non‐host plant species, Vigna unguiculata (Walp.) (Leguminosae) and Vinca rosea (L.) (Apocynaceae). Perhaps, the maxillary palps inform M. sexta mostly about feeding stimulatory chemicals common to most plant species.     

Comparative morphology of sensilla on antenna,maxillary palp and labial palp of larvae of Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) (Coleoptera: Curculionidae)

Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) are two wood boring pests of Ailanthus altissima (Mill.) Swingle (tree of heaven) and the variety Ailanthus altissima var. Qiantouchun. These beetles attack healthy trees and bore into roots and trunks during the larval stage. We studied the typology, distribution and morphostructure of the sensilla on the antennae, maxillary palps and labial palps of E. scrobiculatus and E. brandti larvae using scanning and transmission electron microscopy. The results showed the following: (i) the antennae of the two weevil larvae had two types of sensilla, sensilla basiconica (S.b.1 and S.b.2) and sensilla twig basiconica (S.tb.1‐S.tb.3), with S.tb.4 observed only on the antennae of E. brandti larvae; (ii) the maxillary palps had three types of sensilla, S.b.2, S.tb. (S.tb.2, S.tb.3 and S.tb.5) and digitiform sensilla; (iii) the labial palps had two types of sensilla, S.b.2 and S.tb. (S.tb.2, S.tb.3 and S.tb.5); (iv) the quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant between E. scrobiculatus and E. brandti larvae; and (v) sensilla basiconica had distinct sidewall pores, an apical pore was observed on sensilla twig basiconica, and digitiform sensilla were oval in shape, with a distinct apical pore. Based on the microstructure of the cuticle wall and dendrite, we hypothesized that these sensilla functioned as olfactory, gustatory and hygro‐/thermo‐receptors, respectively. We discuss the relationships among types of sensilla and the types of damage caused by the larvae inside the host tree to understand olfactory and gustatory receptor mechanisms. The results of this study will provide a firm basis for future electrophysiological studies.     

Sensilla of the antennae and the labial and maxillary palps of Blattella germanica (L.) (Dictyoptera: Blattellidae): Their classification and distribution

The various sensilla on the antennae and on the labial and maxillary palps of Blattella germanica (L.) were studied. Thick-walled chemoreceptors with fluted shafts and articulated bases are located on the antennae and on the labial and maxillary palps. Thin-walled chemoreceptors, without fluted shafts or articulated bases, are restricted to the flagellar segments of the antennae and to the distal segments of the palps. Antennae of adult males have more thin-walled chemoreceptors than do those of females. Hair-plate sensilla are found at the scape-head and scape-pedicel joints, and at the joints of segments on the palps. Campaniform sensilla are concentrated as a ring around the distal margin of the pedicel, and are also scattered singly on the scape, pedicel, and flagellar segments of the antennae, and on the first segment of the maxillary palps. Occasionally, a few sensilla coeloconica and cold receptor sensilla are found on the antennal flagellum.     

Antennal responses of Cydia pomonella (L.) exposed to surfaces treated with methoxyfenozide

Electroantennogram (EAG) measurements were recorded from the antennae of male and female codling moth, Cydia pomonella L., to determine whether adult moths exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide experience a decline in their antennal reception and thus olfactory sensitivity. Such a phenomenon would offer a possible mechanism for the previously reported decreased responsiveness from moths treated with methoxyfenozide to pheromone‐ and plant volatile‐based monitoring lures. Mean EAG data revealed that the antennae from methoxyfenozide‐treated male moths appear to be just as sensitive to various doses of synthetic codlemone as the antennae from the control and surfactant‐treated moths, but they appeared to be less sensitive to the pheromone component 12OH (collected from female effluvia) than the control male antennae. Mean male EAG responses to the pheromone components E8,E10‐12Al and codlemone collected from methoxyfenozide‐treated females were significantly less than the responses towards those two pheromone components collected from the control and surfactant‐treated females. Female moth exposure to methoxyfenozide did not negatively impact the sensitivity of female antennae to the plant volatile pear ester, but it did towards the apple volatile butyl hexanoate. Data from this study show that adult C. pomonella exposure to methoxyfenozide‐treated surfaces does appear to negatively impact, in a minor way, the (i) olfactory sensitivity (or detection) of male antennae towards some components of the female sex pheromone, (ii) the female antennal sensitivity towards a key apple volatile and (iii) the attractiveness of female pheromone effluvia.     

Sensing the Underground - Ultrastructure and Function of Sensory Organs in Root-Feeding Melolontha melolontha (Coleoptera: Scarabaeinae) Larvae

Introduction

Below ground orientation in insects relies mainly on olfaction and taste. The economic impact of plant root feeding scarab beetle larvae gave rise to numerous phylogenetic and ecological studies. Detailed knowledge of the sensory capacities of these larvae is nevertheless lacking. Here, we present an atlas of the sensory organs on larval head appendages of Melolontha melolontha. Our ultrastructural and electrophysiological investigations allow annotation of functions to various sensory structures.

Results

Three out of 17 ascertained sensillum types have olfactory, and 7 gustatory function. These sensillum types are unevenly distributed between antennae and palps. The most prominent chemosensory organs are antennal pore plates that in total are innervated by approximately one thousand olfactory sensory neurons grouped into functional units of three-to-four. In contrast, only two olfactory sensory neurons innervate one sensillum basiconicum on each of the palps. Gustatory sensilla chaetica dominate the apices of all head appendages, while only the palps bear thermo-/hygroreceptors. Electrophysiological responses to CO₂, an attractant for many root feeders, are exclusively observed in the antennae. Out of 54 relevant volatile compounds, various alcohols, acids, amines, esters, aldehydes, ketones and monoterpenes elicit responses in antennae and palps. All head appendages are characterized by distinct olfactory response profiles that are even enantiomer specific for some compounds.

Conclusions

Chemosensory capacities in M. melolontha larvae are as highly developed as in many adult insects. We interpret the functional sensory units underneath the antennal pore plates as cryptic sensilla placodea and suggest that these perceive a broad range of secondary plant metabolites together with CO₂. Responses to olfactory stimulation of the labial and maxillary palps indicate that typical contact chemo-sensilla have a dual gustatory and olfactory function.     

Morphological and ultrastructural characterization of olfactory sensilla in Drosophila suzukii: Scanning and transmission electron microscopy

Drosophila suzukii is a serious horticultural and quarantine pest, damaging various berry crops. Although the active use of olfactory communication in D. suzukii is well-known, their olfactory sensory system has not been comprehensively reported. Therefore, the present study was carried out to understand the morphology, distribution and ultrastructure of olfactory sensilla present in the antennae and maxillary palps of D. suzukii, through scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The olfactory sensilla on the antennae of D. suzukii in both sexes could be classified into three major morphological types, basiconic, trichoid and coeloconic sensilla, according to their shapes. The antennal basiconic sensilla were further divided into three subtypes and the antennal trichoid sensilla into two subtypes, respectively, according to the size of individual sensillum. In contrast to the antennal olfactory sensilla showing diverse morphology, basiconic sensilla was the only type of olfactory sensilla in the maxillary palps of D. suzukii. The basiconic sensilla in the maxillary palps could be further classified into three subtypes, based on their size. Our SEM and TEM observations indicated that multiple nanoscale pores are present on the surface of all types of olfactory sensilla in the antennae and maxillary palps, except coeloconic sensilla. The difference in the morphological types and the distribution of olfactory sensilla suggests that their olfactory functions are different between antennae and maxillary palps in D. suzukii. The results of this study provide useful information for further studies to determine the function of olfactory sensilla in D. suzukii and to understand their chemical communication system.     

Effect of food deprivation on attractiveness of food sources, containing natural and artificial sugar and protein, to three African fruit flies: Ceratitis cosyra, Ceratitis fasciventris, and Ceratitis capitata

The effects of food deprivation, age, and mating status on the responses of three fruit fly species, Ceratitis cosyra (Walker), Ceratitits fasciventris (Bezzi), and Ceratitits capitata (Wiedemann) (Diptera: Tephritidae) to natural and artificial sugar and protein food sources were investigated. Natural food sources included guava [Psidium guajava L. (Myrtaceae)] juice (a common host fruit for all three fruit fly species) and bird faeces (farm chicken). Artificial food sources included molasses (obtained from a local sugar factory) and a locally produced protein bait (the International Centre of Insect Physiology and Ecology yeast). In all species studied, sugar deprivation of immature (1–2‐day‐old) male and female flies increased their response to food odours, although it did not change their preference for the type of odour (protein or sugar). Protein deprivation of mature (14–17‐day‐old) male and female flies also increased their response to food odours compared to protein‐fed flies. Protein‐deprived females were highly attracted to odours from protein sources in particular. Odours from natural food sources, guava juice, and chicken faeces, were more attractive to food‐deprived flies than were odours from artificial sugar and protein sources. Attraction to food odours increased significantly with increasing age for protein‐deprived females of all species. For males and females of all species, nutritional state was a more important factor than mating status in influencing responses of flies to food odours. Practical implications of these findings are discussed in terms of strategies for fruit fly control using food baits.     

Background odour induces adaptation and sensitization of olfactory receptors in the antennae of houseflies

The presence of background odour was found to have a small but significant effect on the sensitivity of the antennal olfactory system of houseflies, Musca domestica Linnaeus (Diptera: Muscidae), to new pulses of odour. We show that cross-adaptation and cross-sensitization between a background odour of (+/-)-1-octen-3-ol and pulses of (+/-)-1-octen-3-ol, 2-pentanone and R-(+)-limonene can occur, confirming that olfactory receptor cells are sensitive to different odours. Background odour can increase the responses to low concentration odour pulses and decrease the responses to higher concentration odour pulses. It is suggested that background odour has a larger effect on olfactory receptor cells that respond with a tonic increase of spike frequency, giving information about the level of odour concentration, i.e. the 'static' environment. Cells that respond in a phasic way only provide information on the dynamics of the olfactory environment.     

Extreme sensitivity in an olfactory system

We recorded olfactory-induced cardiac responses to evaluate olfactory response thresholds to behaviourally relevant odours in a moth. Specific antennal receptor neurons enable insects to detect biologically meaningful odours such as sex pheromones and host-plant volatiles. The response threshold values demonstrated here are well below anything earlier reported in any organism. A heart response was triggered by less than six molecules of the most efficient odours hitting the antennae of the insect. The behavioural significance of this extreme sensitivity most likely lies in the creation of awareness and readiness to respond behaviourally at higher concentration levels.     

Premating Behavior of Dealates of the Formosan Subterranean Termite and Evidence for the Presence of a Contact Sex Pheromone

Swarming by alates of the Formosan subterranean termite, Coptotermes formosanus Shiraki, occurs around dusk during April–June in Louisiana and adjoining states. Dispersal is the primary purpose for swarming. We studied the premating behavior, following swarming. Loss of wings, especially in females, was essential for successful formation of tandem pairs between the two sexes. No calling was observed and apparently a long-range volatile sex pheromone, reported in some other species of termites, may not be involved in mate attraction in C. formosanus. Males moving randomly use their antennae to detect a female, and, together with maxillary and labial palps, contact is maintained with her while in tandem. Previously mated males paired with virgin females showed tandem behavior. The incidence was very low when both males and females were previously mated and then paired. Males with complete ablation of antennae did not form tandem pairs. Ablation of 10 terminal antennal segments or the labial palps caused a significant increase in the time to initiate tandem behavior. Electrophysiological recordings from gustatory sensilla on both antennae and maxillary palps revealed increased neural activity in response to female abdominal tip extracts compared to activity elicited by a solvent control. Analysis of extracts of male and female abdominal tips using high-performance liquid chromatography showed a peak unique to the female extract. We discuss the presence of a nonvolatile chemical in female C. formosanus and its involvement in the specific premating behavior.     

The olfactory responses of the antenna and maxillary palp of the fleshfly,Neobellieria bullata (Diptera: Sarcophagidae), and their sensitivity to blockage of nitric oxide synthase

The relative sensitivities of the olfactory receptors in the antenna and maxillary palp of the fleshfly, Neobellieria bullata, were assessed using simultaneous electroantennograms (EAGs) and electropalpograms (EPGs). In general, the antennae and maxillary palps were more sensitive to odors related to animals (blood extract and saturated carboxylic acid) than to odors that were plant-derived (citral, hexenol, hexenal). In addition, the maxillary palps were relatively less sensitive to plant-derived odorants than the antennae, perhaps related to their anatomical position. Scanning electron microscopy was also used to assess the types of sensilla found on the two organs. In addition, NADPH-diaphorase histochemistry was used in an attempt to localize the enzyme nitric oxide synthase (NOS) in the antenna and the maxillary palps. We found evidence of NADPH-diaphorase staining in both organs, with localized staining in the antennal cells and more general staining in the maxillary palps. When NOS was selectively blocked using the antagonist L-NAME, the amplitude of the EAGs and EPGs to odorants fell by 30-50%. In contrast, application of the inactive enantiomer, D-NAME, did not change the amplitude of the EAGs or the EPGs. Our results indicate that NOS is involved in the function of olfactory receptor cells in the fleshfly.     

The composition and timing of flower odour emission by wild <Emphasis Type="Italic">Petunia axillaris</Emphasis> coincide with the antennal perception and nocturnal activity of the pollinator <Emphasis Type="Italic">Manduca sexta</Emphasis>

In the genus Petunia, distinct pollination syndromes may have evolved in association with bee-visitation (P. integrifolia spp.) or hawk moth-visitation (P. axillaris spp). We investigated the extent of congruence between floral fragrance and olfactory perception of the hawk moth Manduca sexta. Hawk moth pollinated P. axillaris releases high levels of several compounds compared to the bee-pollinated P. integrifolia that releases benzaldehyde almost exclusively. The three dominating compounds in P. axillaris were benzaldehyde, benzyl alcohol and methyl benzoate. In P. axillaris, benzenoids showed a circadian rhythm with an emission peak at night, which was absent from P. integrifolia. These characters were highly conserved among different P. axillaris subspecies and P. axillaris accessions, with some differences in fragrance composition. Electroantennogram (EAG) recordings using flower-blends of different wild Petunia species on female M. sexta antennae showed that P. axillaris odours elicited stronger responses than P. integrifolia odours. EAG responses were highest to the three dominating compounds in the P. axillaris flower odours. Further, EAG responses to odour-samples collected from P. axillaris flowers confirmed that odours collected at night evoked stronger responses from M. sexta than odours collected during the day. These results show that timing of odour emissions by P. axillaris is in tune with nocturnal hawk moth activity and that flower-volatile composition is adapted to the antennal perception of these pollinators.     

Single cell recordings from tsetse (Glossina m.morsitans) antennae reveal olfactory, mechano - and cold receptors

Abstract. Action potentials from individual cells and receptor potentials were recorded from antennae (funiculi) of living tsetse flies, Glossina m.morsitans Westwood, using a 'surface-contact' recording technique. Stimuli were pressure of the electrode, changes in temperature and the vapours of l-octen-3–ol, acetone, 3–methylphenol, dichloromethane and CO₂. Two types of mechano-receptive cells were found. One type only fired action potentials when pressure was increased, the other type continued firing when pressure was maintained. Of the 182 cells tested for their sensitivity to temperature change and the odour stimuli, 19% did not respond to any of the stimuli, suggesting that we are still unaware of cues which may be of import to the flies, and 4% responded to temperature only, increasing and decreasing their activity with a decrement and increment in temperature, respectively. Of the 141 cells which were olfactory receptors, 52% responded to 1-octen-3–ol and 40% were exclusively sensitive to this substance. For 3–methylphenol these percentages were 23 and 18, for acetone 13 and 11, for dichloromethane 12 and 7, for CO₂ 13 and 13. No clustering of cell types in certain areas of the funiculi was found, nor was a difference apparent between the sexes. The majority of the olfactory cells responded by increasing their activities on odour stimulation. Inhibition was found in three cells only, in which spontaneous activity was suppressed on stimulation with 3–methylphenol. Spike responses were phasic-tonic and varied between two extremes: Cells showing relatively rapid cessation of spike activity after the end of stimulation, and cells which continued firing for several seconds or minutes after stimulation. Possible behavioural effects of the activities of the various cell types are discussed.     

Behavioural responses of stable flies to cattle manure slurry associated odourants

Stable flies (Stomoxys calcitrans [Diptera: Muscidae] L.) are blood‐feeding synanthropic pests, which cause significant economic losses in livestock. Stable fly antennae contain olfactory sensilla responsive to host and host environment‐associated odours. Field observation indicated that the abundance of stable flies increased significantly in grasslands or crop fields when cattle manure slurry was applied. Major volatile compounds emanating from manure slurry were collected and identified. Behavioural responses of stable flies to those compounds were investigated in laboratory bioassays and field‐trapping studies. Results from olfactometer assays revealed that phenol, p‐cresol and m‐cresol were attractive to adult stable flies. When tested individually, attraction was higher with lower dosages. Stable flies were most attracted to blends of phenol and m‐cresol or p‐cresol. Traps with binary blend lures caught more stable flies in field trials as well.     

Relationship between the olfactory sensilla of Papilio demoleus L. larvae and their orientation responses to different odours

Papilio demoleus larvae showed maximum attraction to odours of citral, cirtonellal and citronellol, slightly less to geraniol and 1-limonene, and low attraction to hydroxy-citronellal and geranyl acetate. These chemicals (except the last) are the main odorous constituents of the insect's host plant (Citrus) leaves. Phellandrene was almost inert, whereas farnesol and d-limonene elicited a low, and linalool, eugenol and clove oil a high repulsion. Larvae deprived of one antenna and both maxillary palps were similar to normal larvae in their orientational responses. Normal larval responses changed with the inactivation of one or more of the three specific antennal sensilla. A full complement of the three sensilla was not essential for the normal larval response to certain chemicals. Any of the sensilla singly or in combination with any other sensillum could mediate the normal larval response (attraction) to citral and geraniol. For a few other chemicals, certain specific sensilla, singly or in a combination of two, mediated the normal response. Only for d-limonene were all three sensilla essential for the normal response.