Antennal lobe projection destinations of Helicoverpa zea male olfactory receptor neurons responsive to heliothine sex pheromone components

We used single sensillum recordings to define male Helicoverpa zea olfactory receptor neuron physiology followed by cobalt staining to trace the axons to destination glomeruli of the antennal lobe. Receptor neurons in type A sensilla that respond to the major pheromone component, (Z)-11-hexadecenal, projected axons to the cumulus of the macroglomerular complex (MGC). In approximately 40% of these sensilla a second receptor neuron was stained that projected consistently to a specific glomerulus residing in a previously unrecognized glomerular complex with six other glomeruli stationed immediately posterior to the MGC. Cobalt staining corroborated by calcium imaging showed that receptor neurons in type C sensilla sensitive to (Z)-9-hexadecenal projected to the dorsomedial posterior glomerulus of the MGC, whereas the co-compartmentalized antagonist-sensitive neurons projected to the dorsomedial anterior glomerulus. We also discovered that the olfactory receptor neurons in type B sensilla exhibit the same axonal projections as those in type C sensilla. Thus, it seems that type B sensilla are anatomically type C with regard to the projection destinations of the two receptor neurons, but physiologically one of the receptor neurons is now unresponsive to everything except (Z)-9-tetradecenal, and the other responds to none of the pheromone-related odorants tested.     

Host plant volatiles synergize responses of sex pheromone-specific olfactory receptor neurons in male <Emphasis Type="Italic">Helicoverpa zea</Emphasis>

Single-cell electrophysiological recordings were obtained from olfactory receptor neurons in antennal trichoid sensilla of male corn earworm, Helicoverpa zea. Spontaneous activity of the neuron specific for the major component ( Z)-11-hexadecenal, the conspecific female-emitted sex pheromone, was not affected by exposure to host plant volatiles. However, stimulations with binary mixtures of a threshold dosage of the pheromone component and increasing dosages of either linalool or ( Z)-3-hexenol significantly synergized the pheromone-specific neuron's firing rates compared with responses to the major pheromone component alone. Cross-adaptation studies confirmed that the enhanced impulses originated from the pheromone-component-tuned neuron. Because plant volatiles do not stimulate the pheromone-specific neuron when presented alone, the pheromone plus host odor blend would be interpreted as containing more pheromone than it actually does when processed by the pheromone-processing portion of the antennal lobe.     

A comparison of responses from olfactory receptor neurons of<Emphasis Type="Italic"> Heliothis subflexa</Emphasis> and<Emphasis Type="Italic"> Heliothis virescens</Emphasis> to components of their sex pheromone

Single-cell electrophysiological recordings were obtained from olfactory receptor neurons in sensilla trichodea on male antennae of the heliothine species Heliothis subflexa and the closely related congener H. virescens. A large percentage of sensilla (72% and 81%, respectively, of all sensilla sampled) contained a single odor-responsive receptor neuron tuned to the major pheromone component of both species, Z-11-hexadecenal. A second population of sensilla on H. subflexa antennae (18%) housed receptor neurons that were tuned to Z-9-hexadecenal but also responded with less sensitivity to Z-9-tetradecenal. A similar population of sensilla (4%) on H. virescens male antennae housed receptor neurons that were shown to be tuned specifically only to Z-9-tetradecenal, with no response to even high dosages of Z-9-hexadecenal. A third population of sensilla (comprising 8% and 16% of the sensilla sampled in H. subflexa and H. virescens, respectively) housed two olfactory receptor neurons, one of which was tuned to Z-11-hexadecenyl acetate and the other tuned to Z-11-hexadecenol. In H. subflexa the Z-11-hexadecenyl acetate-tuned neuron also responded to Z-9-tetradecenal with nearly equivalent sensitivity. The behavioral requirements of males of these two species for distinct pheromonal blends was, therefore, reflected by the subtle differences in the tuning properties of antennal olfactory receptor neurons.Abbreviations MGC macroglomerular complex - ORN olfactory receptor neuron - Z9–14:Ald (Z)-9-tetradecenal - Z9–16:Ald (Z)-9-hexadecenal - Z11–16:Ac (Z)-11-hexadecenyl acetate - Z11–16:Ald (Z)-11-hexadecenal - Z11–16:OH (Z)-11-hexadecenol     

The macroglomerular complex of the antennal lobe in the tobacco budworm moth Heliothis virescens : specified subdivision in four compartments according to information about biologically significant compounds

The functional organisation of the male specific macroglomerular complex in Heliothis virescens has been studied by tip recordings of sensilla trichodea type 1 combined with cobalt-lysine stainings and by intracellular recordings of antennal lobe projection neurons combined with neurobiotin stainings. The antennal lobe, the macroglomerular complex and the stained axons/dendrites were reconstructed by camera-lucida. Some were further computer reconstructed in three dimensions. The results showed that: 1) The macroglomerular complex consisted of four anatomically separated compartments; 2) A large compartment (the cumulus) at the entrance of the antennal nerve received input from receptor neurons responding to the major pheromone component; 3) Another large compartment, located dorso-medially of the cumulus (the dorso-medial compartment) received input from receptor neurons tuned to the second pheromone component; 4) Two ventrally located compartments received input from two receptor neuron types, co-localized in the same sensillum. Each neuron type responded strongest to one of two interspecifically acting signals, shown to interrupt the pheromone attraction. 5) The function of the dorso-medial compartment was further verified by selective arborizations in this compartment by a projection neuron showing strong response to antennal stimulation with the second pheromone component. At low concentration, the neuron responded synergistically to stimulation with the binary pheromone mixture. Accepted: 29 July 1998     

Incomplete electrical isolation of sex-pheromone responsive olfactory receptor neurons from neighboring sensilla

In the long trichoid sensilla on male Helicoverpa zea antennae, approximately 40% of the sensilla having a large-spiking olfactory receptor neuron responding to the major pheromone component, (Z)-11-hexadecenal, also exhibit small-spiking action potentials that also seem to be responsive to this same compound. In this study, we investigated whether these small-spiking signals are a result of intrusive electrical signals generated from neighboring sensilla. Two methods were used for this study. First, the sensillum was completely covered by the saline-filled recording electrode to physically prevent the sensillum from being contacted by exposure to (Z)-11-hexadecenal. In this case, activation of the large-spiking neuron in response to the pheromone component was prevented, whereas the small-spiking activity continued to be influenced by the airborne delivery of the pheromone. In the second method the (Z)-11-hexadecenal was applied directly in solution through the cut tip of the sensillum through the recording electrode. In this case only large-spiking activity occurred in response to (Z)-11-hexadecenal, with no increase whatsoever in the firing frequency of the small spikes. We conclude that these long trichoid olfactory sensilla are not completely isolated electrically from neighboring sensilla and that small spikes in some recordings originate from large-spiking olfactory receptor neurons (ORNs) in neighboring sensilla.     

Functional map of the macroglomerular complex of male Helicoverpa armigera

The mechanism of sex pheromone reception in the male cotton bollworm Helicoverpa armigera has been extensively studied because it has become an important model system for understanding insect olfaction. However, the pathways of pheromone processing from the antenna to the primary olfactory center in H. armigera have not yet been clarified. Here, the physiology and morphology of male H. armigera olfactory sensory neurons (OSNs) were studied using single sensillum recording along with anterograde filling and intracellular recording with retrograde filling. OSNs localized in type A sensilla responded to the major pheromone component cis-11-hexadecenal, and the axonal terminals projected to the cumulus (Cu) of the macroglomerular complex (MGC). The OSNs in type B sensilla responded to the behavioral antagonist cis-9-tetradecenal, and the axonal terminals projected to the dorsomedial anterior (DMA) unit of the MGC. In type C sensilla, there were 2 OSNs: one that responded to cis-9-tetradecenal and cis-11-hexadecenol with the axonal terminals projecting to the DMA, and another that responded to the secondary pheromone components cis-9-hexadecenal and cis-9-tetradecenal with the axonal terminals projecting to the dorsomedial posterior (DMP) unit of the MGC. Type A and type B sensilla also housed the secondary OSNs, which were silent neurons with axonal terminals projected to the glomerulus G49 and DMP. Overall, the neural pathways that carry information on attractiveness and aversiveness in response to female pheromone components in H. armigera exhibit distinct projections to the MGC units.     

棉铃虫雄蛾触角的毛形感器对其性信息素组分及类似物的反应

本文用单细胞电反应的记录方法,测定了棉铃虫Heliothis armigera Hubner雄蛾触角的毛形感器对其性信息素组分及雌蛾腹部提取物的反应,发现顺-11-十六碳烯醛和顺-9-十六碳烯醛能引起反应。对前者发放大脉冲,对后者发放小脉冲,对雌蛾腹部提取物发放大、小两种脉冲,但以大脉冲为主。     

Functional divergence of spatially conserved olfactory glomeruli in two related moth species

In different moth species, the number and spatial arrangement of olfactory glomeruli in the antennal lobe (AL) vary widely, but the spatial map within a species is thought to be invariant, making it possible to identify single glomeruli across individuals. We investigated the relationship between the physiological tuning of pheromone-selective interneurons and their association with specific, identified glomeruli in the macroglomerular complex (MGC) of the noctuid moth, Heliothis subflexa. Three odorants that are required for pheromone-source location in this species were tested individually and in blends. Recordings from 27 pheromone-specific projection neurons (PNs) indicated that the majority (48%) were selectively activated by the major pheromone component of this species, Z-11-hexadecenal (Z11-16:Ald), with 33% primarily tuned to Z-9-hexadecenal and 19% to Z-11-hexadecenol. Intracellular staining revealed that the dendrites of PNs tuned to Z11-16:Ald always branched within the largest glomerulus of the MGC, the cumulus. Similarly, each of the other two classes of PN was associated with a different 'satellite' glomerulus in the MGC. The spatial configuration of the four-glomerulus H. subflexa MGC was indistinguishable from that previously reported in the closely related species, Heliothis virescens. Hence, as these two species diverged, changes in the association of satellite MGC glomeruli with particular odorants have occurred without a measurable change in the anatomical arrangement of the glomerular array.     

雄性棉铃虫感受性信息素的分子和神经机制研究进展

棉铃虫Helicoverpa armigera主要借助于性信息素通讯完成雌雄识别,实现交配和种群繁衍。关于棉铃虫感受性信息素机制的研究一直是我国化学生态学领域的热点和重心,研究结果有助于开发和改进棉铃虫防治的性引诱剂。本文将对棉铃虫雄虫感受雌虫释放的性信息素的机制进行综述,以期为深入研究棉铃虫及其他相关昆虫的性信息素感受的分子和神经机理提供参考。棉铃虫雌虫性信息素腺体合成和释放多种长链、饱和或非饱和的脂肪醛和醇等化合物,其中Z11-16:Ald为主要性信息素成分,Z9-16∶Ald和Z9-14∶Ald为次要性信息素成分,不同组分按一定比例混合可明显增强对雄性棉铃虫的引诱效果,而化合物Z11-16∶OH和高剂量的Z9-14∶Ald对性信息素引诱活性具有明显的抑制效果。相应地,雄性棉铃虫触角上A, B和C 3种类型的毛形感器能够感受这些信息化合物。A类型毛形感器内表达受体OR13感受Z11-16∶Ald,B类型毛形感器内表达OR14b感受Z9-14∶Ald,C类型毛形感器内表达OR6和OR16感受Z9-16∶Ald, Z9-14∶Ald, Z11-16:Ac和Z11-16∶OH。受体的表达位置和功能与不同类型毛形感器的电生理反应特性相一致。钙离子成像证明在棉铃虫触角叶内的3个扩大型神经纤维球接受这些气味信息,其中神经纤维球云状体接受Z11-16∶Ald,背中间后侧纤维球接受Z9-16∶Ald,背中间前侧纤维球接受Z9-14∶Ald, Z11-16∶Ac和Z11-16∶OH。这些研究成果在感器、受体和脑中枢水平上揭示了棉铃虫感受性信息素的机制,在这些研究基础上,我们认为需要深入开展以下方面的研究：(1)进一步鉴定相关性信息素受体的功能和定位;(2)深入研究脑内嗅觉高级中枢对性信息素信息的处理和整合神经机制;(3)明确棉铃虫性信息素感受受到寄主植物、光周期、温度、湿度等环境因素的影响及机制。     

Functional organization of the macroglomerular complex related to behaviourally expressed olfactory redundancy in male cabbage looper moths

Abstract. The neurophysiological bases for behaviourally expressed olfactory redundancy in the sex pheromone communication system of the cabbage looper moth, Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae), were examined by coupling the cut-sensillum extracellular recording technique with a highly specific neuronal marking method for moth peripheral receptors. In seventy-two antennal sensilla, axonal pathways of cobalt-stained neurones could be traced into the male-specific macroglomerular complex in the antennal lobe. In T. ni males this comprises five glomeruli, two of which are subdivided into morphologically, and in some instances functionally identifiable, regions. Axonal arborizations of forty-eight neurones (single stainings) showed high fidelity (98%) for containment within a specific glomerulus or glomerular subdivision, and the neuropil targeted seemed to be related to the specificity of a neurone to a particular female-emitted sex pheromone component (27-12:Ac, Z7-14:Ac, Z9-14:Ac, 12:Ac, 11–12:Ac, Z5-12:Ac), or to a behavioural antagonist (Z7-12:OH). Double (twenty-one) and multiple stainings (three) showed axons projecting into two or more glomeruli, respectively, with 100% fidelity for the component-specific glomerulus or glomerular subdivision to be targeted. We suggest that the potential for a single minor component to cross-stimulate two or more neurones within a sensillum may enable partial blends to continue to provide sensory input into all of the pheromone-processing glomeruli of the complex. Our interpretation is that redundancy occurs at the receptor level on neighbouring dendrites, and thus allows various four-component partial blends to evoke full pheromone-mediated behaviour.     

Distinct Projections of Two Populations of Olfactory Receptor Axons in the Antennal Lobe of the Sphinx Moth Manduca sexta

The central projections of olfactory receptor cells associatedwith two distinct types of antennal sensilla in the sphinx mothManduca sexta were revealed by anterograde staining. In bothsexes, receptor axons that arise from sexually isomorphic, type-IItrichoid sensilla (and possibly some basiconic sensilla) projectto the spheroidal glomeruli in the ipsilateral antennal lobe.Each axon terminates in one glomerulus. Axons from a limitedregion of the antenna project to glomeruli throughout the lobe,arguing against strict topographic mapping of antennal receptorcells onto the array of glomeruli. Axons of sex-pheromone-selectivereceptor cells in the male-specific type-I trichoid sensillaproject exclusively to the sexually dimorphic macroglomerularcomplex (MGC). Axons from sensilla on the dorsal surface ofthe antenna are biased toward the medial MGC and those fromventral sensilla, toward the lateral MGC. Some receptor-cellaxons branch before reaching the MGC, but their terminals arealways confined to one of the two main glomerular divisionsof the MGC, the cumulus and toroid. These findings confirm thatprimary-afferent information about pheromonal and non-pheromonalodors is segregated in the antennal lobe and suggest that thereis a functional correspondence between particular olfactoryreceptor cells and specific glomeruli. Chem. Senses 20: 313–323,1995.     

Physiology and glomerular projections of olfactory receptor neurons on the antenna of female Heliothis virescens (Lepidoptera: Noctuidae) responsive to behaviorally relevant odors

The neurophysiology and antennal lobe projections of olfactory receptor neurons housed within short trichoid sensilla of female Heliothis virescens F. (Noctuidae: Lepidoptera) were investigated using a combination of cut-sensillum recording and cobalt-lysine staining techniques. Behaviorally relevant odorants, including intra- and inter-sexual pheromonal compounds, plant and floral volatiles were selected for testing sensillar responses. A total of 184 sensilla were categorized into 25 possible sensillar types based on odor responses and sensitivity. Sensilla exhibited both narrow (responding to few odors) and broad (responding to many odors) response spectra. Sixty-six percent of the sensilla identified were stimulated by conspecific odors; in particular, major components of the male H. virescens hairpencil pheromone (hexadecanyl acetate and octadecanyl acetate) and a minor component of the female sex pheromone, (Z)-9-tetradecenal. Following characterization of the responses, olfactory receptor neurons within individual sensilla were stained with cobalt lysine (N=39) and traced to individual glomeruli in the antennal lobe. Olfactory receptor neurons with specific responses to (Z)-9-tetradecenal, a female H. virescens sex pheromone component, projected to the female-specific central large female glomerulus (cLFG) and other glomeruli. Terminal arborizations from sensillar types containing olfactory receptor neurons sensitive to male hairpencil components and plant volatiles were also localized to distinct glomerular locations. This information provides insight into the representation of behaviorally relevant odorants in the female moth olfactory system. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

利用单感器记录与神经元示踪结合对棉铃虫主要性信息素感器内神经元投射的鉴定

【目的】鉴定雄性棉铃虫Helicoverpa armigera成虫触角性信息素感器嗅觉受体神经元的功能、形态及中枢投射路径。【方法】利用单感器记录技术记录棉铃虫嗅觉受体神经元对性信息素的反应,同时采用荧光染料作为示踪剂染色标记嗅觉受体神经元;使用免疫组织化学方法处理相应的脑组织,标记脑内触角叶的神经纤维球结构;用激光扫描共聚焦显微镜获取图像数据,使用图形软件ZEN和Amira 4.1.1进行三维结构重建。【结果】记录到雄性棉铃虫成虫触角上长毛形感器对主要性信息素成分Z11-16∶Ald产生明显的电生理反应,并成功染色标记了该感器内的嗅觉受体神经元。染色标记显示该感器内具有两个嗅觉受体神经元,其轴突通过触角神经分别投射触角叶内的云状体神经纤维球和普通神经纤维球。【结论】单感器记录与神经元示踪两技术结合能够用于鉴定昆虫触角嗅觉受体神经元的功能、形态和投射至神经纤维球的路径。与赖氨酸钴方法比较,使用荧光染料法进行神经元示踪,操作更简便,且易于进行三维空间分析,为调查棉铃虫其他嗅觉神经元的投射路径以明确外周气味受体感受与中枢系统的联系提供了有力技术支持。     

Neurons discovered in male Helicoverpa zea antennae that correlate with pheromone-mediated attraction and interspecific antagonism

Responses of single receptor neurons in the antennae of male Helicoverpa zea to sex pheromone components and to behavioral antagonists were recorded using a cut-sensillum extracellular recording technique. Three types of sensilla were identified from sampling 325 male-specific sensilla trichodea located at the lateral edge of antennomeres. The majority of these sensilla (71%) contained a receptor neuron tuned to the principal sex pheromone component (Z)-11-hexadecenal. A second sensillar type (10%) contained a receptor neuron that responded only to (Z)-9-tetradecenal. A third sensillar type (19%) contained a large-spiking neuron tuned to the secondary pheromone component (Z)-9-hexadecenal, but this neuron also could be stimulated to equivalent spike frequencies by the same emitted amounts of (Z)-9-tetradecenal. A smaller-spiking neuron in this sensillar type responded to two compounds known to act only as behavioral antagonists, (Z)-11-hexadecen-1-ol and (Z)-11-hexadecenyl acetate, and to (Z)-9-tetradecenal. Cross-adaptation studies confirmed the presence of one large- and one small-spiking neuron in the third sensillar type. Dose-response studies correlated to collected stimuli amounts showed that the large-spiking neuron in the third sensillar type was equally tuned to (Z)-9-hexadecenal and (Z)-9-tetradecenal, whereas the smaller-spiking neuron was far more sensitive to (Z)-11-hexadecen-1-ol and to (Z)-11-hexadecenyl acetate than to (Z)-9-tetradecenal. Accepted: 29 September 1997     

Responses of a population of antennal olfactory receptor cells in the female moth Manduca sexta to plant-associated volatile organic compounds

Extracellular electrophysiological recordings were made from individual type-A trichoid sensilla on the antenna of the female sphinx moth Manduca sexta. A single annulus of the antenna bears about 1,100 of these sensilla, and each is innervated by two olfactory receptor cells. We tested the responses of these receptor cells to a panel of 102 volatile compounds, as well as three plant-derived odor mixtures, and could discern three different functional types of type-A trichoid sensilla. One subset of receptor cells exhibited an apparently narrow molecular receptive range, responding strongly to only one or two terpenoid odorants. The second subset was activated exclusively by aromatics and responded strongly to two to seven odorants. The third subset had a broad molecular receptive range and responded strongly to odorants belonging to several chemical classes. We also found receptor cells that did not respond to any of the odorants tested but were spontaneously active. Certain odorants elicited excitatory responses in some sensilla but inhibitory responses in others, and some receptor cells were strongly excited by certain odorants but inhibited by others. Impregnation of groups of receptor cells in type-A trichoid sensilla with rhodamine-dextran demonstrated that their axons project mainly to the large female glomeruli of the antennal lobe.     

Responses of a population of antennal olfactory receptor cells in the female moth Manduca sexta to plant-associated volatile organic compounds.

Extracellular electrophysiological recordings were made from individual type-A trichoid sensilla on the antenna of the female sphinx moth Manduca sexta. A single annulus of the antenna bears about 1,100 of these sensilla, and each is innervated by two olfactory receptor cells. We tested the responses of these receptor cells to a panel of 102 volatile compounds, as well as three plant-derived odor mixtures, and could discern three different functional types of type-A trichoid sensilla. One subset of receptor cells exhibited an apparently narrow molecular receptive range, responding strongly to only one or two terpenoid odorants. The second subset was activated exclusively by aromatics and responded strongly to two to seven odorants. The third subset had a broad molecular receptive range and responded strongly to odorants belonging to several chemical classes. We also found receptor cells that did not respond to any of the odorants tested but were spontaneously active. Certain odorants elicited excitatory responses in some sensilla but inhibitory responses in others, and some receptor cells were strongly excited by certain odorants but inhibited by others. Impregnation of groups of receptor cells in type-A trichoid sensilla with rhodamine-dextran demonstrated that their axons project mainly to the large female glomeruli of the antennal lobe.     

Chemical communication in heliothine moths

The chemical and temporal features of the sex-pheromone emitted by Heliothis virescens females are encoded by a diverse array of output pathways from the male-specific macroglomerular complex (MGC) in the antennal lobe. Most output neurons (29 out of 32) were activated by antennal stimulation with the principal component of the sex-pheromone blend of this species, (Z)-11-hexadecenal. Six neurons were excited solely by this component, 8 neurons also responded to the second essential blend component, (Z)-9-tetradecenal, and 14 neurons displayed equivalent responses to the two. Many neurons also effectively encoded the onset and duration of the stimulus. In one additional neuron, a prolonged excitatory response (synergism) was evoked only by the blend of the two components, indicating that some MGC neurons function as blend detectors.In contrast to the situation in Helicoverpa zea, none of the MGC neurons in H. virescens responded selectively to (Z)-9-tetradecenal, suggesting that these two noctuid species employ different neural strategies to encode information about their respective pheromone blends.Three MGC-output neurons responded selectively to (Z)-11-hexadecenyl acetate, an odorant released by some sympatric species that disrupts normal upwind flight to pheromones. Thus, changes in the attractant and deterrent chemical signals, as well as the physical features of these odor plumes, are encoded in the MGC across a diverse parallel array of output pathways to the protocerebrum.Abbreviations AL antennal lobe - AN antennal nerve - 16:AL hexadecanal - MGC macroglomerular complex - 14:AL tetradecanal - Z11-16:AL (Z) 11-hexadecenal - Z11-16:AC (Z) 11-hexadecenyl acetate - Z9-14:AL (Z) 9-tetradecenal - Z9-14:FO (Z) 9-tetradecenyl formate     

Detection of sex pheromone components in Manduca sexta (L.).

The ability of olfactory receptor neurons to detect female-produced sex pheromone components and a limited sample of potential host plant odours was studied by single-sensillum recordings from olfactory sensilla present on male and female antennae in Manduca sexta. The majority of pheromone-sensitive receptor neurons examined in males was specialized for detection of the two major pheromone components, E10,Z12-hexadecadienal and E10,E12,Z14-hexadecatrienal or E10,E12,E14-hexadecatrienal. New olfactory receptor neurons tuned to the minor components E10,E12-hexadecadienal and Z11-hexadecenal were found. In females, olfactory receptor neurons specific to Z11-hexadecanal were discovered. Pheromone components and host volatiles were detected by separate sets of receptor neurons.     

Antennal neurones specific for redundant pheromone components in normal and mutant Trichoplusia ni males

Abstract Recordings were made from the pheromone-sensitive receptor cells within antennal hairs of normal and mutant male cabbage loopers, Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae), using a cut-sensillum technique. From sampling 136 sensilla on normal males and 123 on mutant males, cells excited by pairs of behaviourally redundant minor pheromone components were discovered: Z9–14: Ac was found to be replaceable with 12: Ac and 11–12: Ac was found to be replaceable with Z5–12: Ac. These cells were not found during previous neurophysiological investigations, but explain most of the associations between mutually replaceable (redundant) pheromone components which had been demonstrated previously to be behaviourally redundant in wind tunnel studies. Our results indicate that the mutant gene in T.ni that affects pheromone production does not affect pheromone receptors in males. Using both AC- and DC-coupled recordings from receptor cells, we found that a single minor component could apparently hyperpolarize one cell while depolarizing another cell within the same sensillum, suggesting that noise reduction and other complex signal processing by receptor cells may contribute to odour processing in the macroglomerulus of the antennal lobe.     

Response specificity of male olfactory receptor neurones for the major and minor components of a female pheromone blend

Abstract The sex attractant of the female redbanded leafroller moth, Argyrotaenia velutinana (Walker), is a blend of seven compounds. Specialized olfactory receptor neurones had been found for only two of the compounds, (Z)-11-tetradecenyl acetate (Z11-14:Ac) and (E)-11-tetradecenyl acetate (E11–14:Ac). These receptor neurones were always found in pairs within the long trichoid sensilla, which are the most abundant multi-pored sensilla on the male antenna. A systematic survey of all regions of the male antenna with standard extracellular recording techniques was undertaken to find receptor neurones responsive to the remaining five minor components of the female pheromone. Of the 113 long trichoid sensilla sampled, all contained two receptor neurones, one specialized for Z11–14:Ac and a second specialized for Ell –14:Ac. A comparable number of recordings were then obtained from the less abundant classes of multi-pored sensilla. Two new receptor neurone types were found, responsive to the minor pheromone components (E)-9-dodecenyl acetate (E9-12:Ac) and (Z)-9-dodecenyl acetate (Z9-12:Ac). Scanning electron micrographs indicated that these recordings were obtained from shorter, narrower trichoid sensilla. The majority of these sensilla appeared to contain three neurones capable of spontaneous action potential production. In each sensillum, only one receptor neurone appeared to respond to stimulation with a minor component of the female blend. The remaining two neurones did not respond to the chemical stimuli evaluated.