Different seasonal rearing conditions do not affect pheromone-sensitive receptor neurons of the adult cabbage looper moth, Trichoplusia ni

Abstract. In southern parts of the United States the cabbage looper moth, Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae), is multivoltine, and therefore successive generations experience different environmental conditions during development from larvae to adults.Since environmental conditions are thought to influence pheromone communication, we tested the effects of two different temperature and light regimes (selected to mimic those occurring in the spring and summer growing seasons in the south) during rearing on the response characteristics of the adult male olfactory receptor neurons responsible for detecting the major component of the female pheromone.The dose-response functions of receptor neurons from the warm- and cold-reared insects were similar in both their slopes and thresholds to stimulation with the major component of the female-emitted pheromone, (Z)-7, dodecen-1-ol acetate and (Z)-7, dodecen-1-ol, a behavioural inhibitor.In double pulse experiments, designed to emulate the temporal dispersion of pheromone in nature, neurons were stimulated with short pulses (200 ms) of (Z)-7, dodecen-1-ol acetate separated by varying intervals.Intervals as short as 30 ms reduced the response to a second pulse by over 50%.When the intervals between pulses were longer than 2 s, significant differences were not seen between the responses to the first and to the second pulse.These temporal response patterns were similar in both warm- and cold-reared animals.     

A general odorant background affects the coding of pheromone stimulus intermittency in specialist olfactory receptor neurones

In nature the aerial trace of pheromone used by male moths to find a female appears as a train of discontinuous pulses separated by gaps among a complex odorant background constituted of plant volatiles. We investigated the effect of such background odor on behavior and coding of temporal parameters of pheromone pulse trains in the pheromone olfactory receptor neurons of Spodoptera littoralis. Effects of linalool background were tested by measuring walking behavior towards a source of pheromone. While velocity and orientation index did drop when linalool was turned on, both parameters recovered back to pre-background values after 40 s with linalool still present. Photo-ionization detector was used to characterize pulse delivery by our stimulator. The photo-ionization detector signal reached 71% of maximum amplitude at 50 ms pulses and followed the stimulus period at repetition rates up to 10 pulses/s. However, at high pulse rates the concentration of the odorant did not return to base level during inter-pulse intervals. Linalool decreased the intensity and shortened the response of receptor neurons to pulses. High contrast (>10 dB) in firing rate between pulses and inter-pulse intervals was observed for 1 and 4 pulses/s, both with and without background. Significantly more neurons followed the 4 pulses/s pattern when delivered over linalool; at the same time the information content was preserved almost to the control values. Rapid recovery of behavior shows that change of perceived intensity is more important than absolute stimulus intensity. While decreasing the response intensity, background odor preserved the temporal parameters of the specific signal.     

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.     

Temporal resolution of odour pulses by three types of pheromone receptor cells inAntheraea polyphemus

Summary Temporal response characteristics of three cell types of maleA. polyphemus, each responding to a different pheromone component, have been measured using series of short (20 ms) pheromone pulses. The stimuli were delivered through capillaries 20 m in diameter and applied to single olfactory sensilla trichodea. Two of three cell types sensitive to (E,Z)-6,11-hexadecadienal and (E,Z)-4,9-tetradecadienyl acetate are able to resolve at least 5 stimuli/s whereas the third, responding to the major pheromone component (E,Z)-6,11-hexadecadienyl acetate, is slower, resolving only about 2 stimuli/s. These results suggest that receptor cells are able to respond to pulses of pheromone concentration as they occur downwind from a point source. The time-averaged number of nerve impulses does not seem to be a reliable measure of the amount of pheromone reaching the sensillum. Responses of the cells thus reflect the non-uniform distribution of pheromone in a plume rather than the average concentration.     

A comparative study of pheromone perception in two species of Noctuid moths

The electrical activity of single olfactory receptor neurons in male soybean looper (SBL) Pseudoplusia includens(Walker) and cabbage looper (CL) Trihoplusia ni(Hübner) moths was evaluated in response to stimulation with fixed amounts of the individual components of their respective pheromone blends. In common with earlier observations in the CL, there are at least two classes of morphologically distinct pheromone sensitive sensilla on the antenna of male SBL, each of which contains two olfactory receptor neurons. In both species, one class of sensilla contains an olfactory receptor neuron sensitive to (Z)-7-dodecen-1-ol acetate (Z-7, 12:AC), the major component in each insect's blend, and a companion receptor neuron which is sensitive to (Z)-7-dodecen-1-ol (Z7,12: OH). In both species the second class of sensilla contains an olfactory receptor neuron which is sensitive to one of the minor components of the pheromone blend. (Z)-5-dodecen-1-ol acetate (Z-5,12:AC) is an effective stimulus in SBL, whereas (Z)-7-tetradecen-1-ol acetate (Z-7,14:AC) is an effective stimulus in CL. However, these two stimulatory compounds have been identified only in the female CL gland; neither has been found in the SBL gland. Thus, in contrast to the CL, which has receptor neurons which are responsive exclusively to conspecific pheromone components, the SBL has a class of receptor neurons which is responsive to a minor component of another species' pheromone blend. Field-trapping assays in which Z-5,12:AC is added to the SBL blend suggest that this single CL component is a powerful inhibitor of male SBL behavioral responses to conspecific pheromone blends. The difference observed in the specificity of the receptor neurons in this second class of sensilla are thus believed to play an integral role in the isolation processes that are maintained between these two species and may well account for the observed behavioral differences in their responses to heterospecific pheromone blends.     

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.     

Glomerular targets of Heliothis subflexa male olfactory receptor neurons housed within long trichoid sensilla

We used single-sensillum recordings to characterize male Heliothis subflexa antennal olfactory receptor neuron physiology in response to compounds related to their sex pheromone. The recordings were then followed by cobalt staining in order to trace the neurons' axons to their glomerular destinations in the antennal lobe. Receptor neurons responding to the major pheromone component, (Z)-11-hexadecenal, in the first type of sensillum, type-A, projected axons to the cumulus of the macroglomerular complex (MGC). In approximately 40% of the type-A sensilla, a colocalized receptor neuron was stained that projected consistently to the posterior complex 1 (PCx1), a specific glomerulus in an 8-glomerulus complex that we call the Posterior Complex (PCx). We found that receptor neurons residing in type-B sensilla and responding to a secondary pheromone component, (Z)-9-hexadecenal, send their axons to the dorsal medial glomerulus of the MGC. As in the type-A sensilla, we found a cocompartmentalized neuron within type-B sensilla that sends its axon to a different glomerulus of the PCx4. One neuron in type-C sensilla tuned to a third pheromone component, (Z)-11-hexadecenol, and a colocalized neuron responding to (Z)-11-hexadecenyl acetate projected their axons to the anteromedial and ventromedial glomeruli of the MGC, respectively.     

Neurophysiological responses of pheromone-sensitive receptor neurons on the antenna of Trichoplusia ni (Hubner) to pulsed and continuous stimulation regimens

Both the frequency and the temporal pattern of action potentialproduction in an insect olfactory receptor neuron are stronglyaffected by odorant composition and the time course over whichstimulus concentration varies. To investigate the temporal characteristicsof the neurophysiological responses of these neurons, we deviseda stimulus delivery system that allows us to repeatedly presentwell-mixed, constant concentration odor pulses with relativelysharp onsets and offsets. Here we compare neurophysiologicalresponses to several different stimulation regimens, includingpulses of different durations and repetition rates. During stimulationwith high concentrations of pheromone, the temporal patternof neural activity from olfactory receptor neurons on the antennaof Trichoplusia ni (Hübner) is characterized by an initialphasic period (100–200 ms), followed by a tonic periodwhich is typically maintained for the remaining duration ofthe stimulus. Different olfactory receptor neurons appear tovary among themselves in the relative distribution between thephasic and tonic portions of the overall discharge. During stimulationregimens involving rapid repeated pulses of odorants, a portionof the phasic response levels is preserved during each pulse.Consequently, T. ni males probably detect much of the fluctuationin concentration of pheromone that may normally occur downwindfrom the site of pheromone release.     

Sound production in the whitemouth croaker and relationship between fish size and disturbance call characteristics

The whitemouth croaker produces two different sounds using extrinsic sonic muscles: (1) male advertisement calls during the spawning season and (2) disturbance calls, produced by both sexes. The advertisement call, related to courtship, was recorded in the field and from two marked spawning males of 28 and 30.5 cm L_T in the laboratory. It consists of individual pulses with average durations of 19.7 ms and 17.8 ms for the two males respectively, interpulse intervals of 496 ms and 718 ms, and dominant frequencies of 280 Hz and 316 Hz. Pulses are emitted in bouts of one to three min duration. Disturbance calls consist of a burst of pulses produced at short intervals, and the pulse duration, interpulse interval and dominant frequency of the pulses average 19.8 ms, 17.1 ms, and 363 Hz, respectively. Dominant frequency and interpulse interval decrease and pulse duration increases with fish size. Sound characteristics change markedly in young of the year individuals (lower than 25 cm L_T) after which they appear to stabilize. Higher dominant frequency in the advertisement than in the disturbance call and the relationship of dominant frequency to pulse duration suggest that dominant frequency is determined as a forced response to muscle contraction parameters rather than by the natural frequency of the swimbladder.     

Biogenic amines modulate olfactory receptor neurons firing activity in Mamestra brassicae

The modulatory effects of the biogenic amines octopamine and serotonin on pheromonal receptor neurons of Mamestra brassicae were investigated. The responses to sex pheromone components of two cells types (A and B) in single male long sensilla trichodea were monitored. Cell types A and B do not respond to the same compound. The response of type A to a pulse of the major sex pheromone component increased 5 min after octopamine injection. Responses of type B to other odorants increased after 30 min. In the absence of any pheromone stimulation the background firing activity of type A increased following octopamine injection. This background activity was used to evaluate the kinetics of octopamine and other biogenic amine effects on olfactory receptor neurons. Octopamine increased this background activity in a concentration- and time-dependent manner. Clonidine, an octopamine agonist, was shown to be more powerful in increasing the background activity of olfactory receptor neurons. The effects of octopamine and clonidine were hypothesized to arise from specific receptor activation as chlorpromazine (an octopamine antagonist) was shown to block the effect of octopamine. Serotonin, a known neuromodulator in most animal species, induced a reversible inhibition of spike firing. Altogether, these results indicate that biogenic amines can modulate the sensitivity of olfactory receptor neurons of moths either directly or by an action on adaptation.     

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.     

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.     

Volatile organic compounds as signals in a plant-herbivore system: electrophysiological responses in olfactory sensilla of the moth Cactoblastis cactorum

The morphological sensillum types on the antennae of male and female Cactoblastis cactorum were visualized by scanning electron microscopy. Electrophysiological recordings were performed for the first time on single olfactory sensilla of C. cactorum. The male sensilla trichodea house a receptor cell responding to the putative pheromone component (9Z,12E)-tetradecadienyl acetate. The sensilla trichodea of the females were much shorter than those of the males and contained specialized receptor cells responding to certain terpenoids, the most frequent being the nerolidol-sensitive cell. The sensilla auricillica and sensilla basiconica of both sexes contained cells responding less specifically to terpenoid compounds as well as to green leaf volatiles. Cells of the sensilla coeloconica responded to aliphatic aldehydes and acids. Eight volatile organic compounds emitted by Opuntia stricta, a host plant of C. cactorum, were identified using gas chromatography-mass spectrometry, beta-caryophyllene being the major compound. Five compounds identified by gas chromatography in the headspace of O. stricta elicited responses in olfactory receptor cells of C. cactorum, nonanal being the most active compound and therefore a candidate attractant of C. cactorum.     

Extracellular transduction events under pulsed stimulation in moth olfactory sensilla

In natural conditions, pheromones released continuously by female moths are broken in discontinuous clumps and filaments. These discontinuities are perceived by flying male moths as periodic variations in the concentration of the stimulus, which have been shown to be essential for location of females. We study analytically and numerically the evolution in time of the activated pheromone-receptor (signaling) complex in response to periodic pulses of pheromone. The 13-reaction model considered takes into account the transport of pheromone molecules by pheromone binding proteins (PBP), their enzymatic deactivation in the perireceptor space and their interaction with receptors at the dendritic membrane of neurons in Antheraea polyphemus sensitive to the main pheromone component. The time-averaged and periodic properties of the temporal evolution of the signaling complex are presented, in both transient and steady states. The same time-averaged response is shown to result from many different pulse trains and to depend hyperbolically on the time-averaged pheromone concentration in air. The dependency of the amplitude of the oscillations of the signaling complex on pulse characteristics, especially frequency, suggests that the model can account for the ability of the studied type of neuron to resolve repetitive pulses up to 2 Hz, as experimentally observed. Modifications of the model for resolving pulses up to 10 Hz, as found in other neuron types sensitive to the minor pheromone components, are discussed.     

Responses to sex pheromone and plant odours by olfactory receptor neurons housed in sensilla auricillica of the codling moth, Cydia pomonella (Lepidoptera: Tortricidae)

Antennal olfactory receptor neurons located in a limited number of two types of sensilla auricillica, the rabbit-eared shoehorn and the regular shoehorn, located on the 5-30 flagellomere of the codling moth, Cydia pomonella, antenna were screened for selectivity to 11 plant compounds, the major sex pheromone component, three minor pheromone components and one behavioural antagonist. Both types of sensilla housed at least three neurons characterised by different action potential amplitudes. Neurons in both males and females responded to the plant compounds, ethyl (E,Z)-2,4-decadienoate, (+/-)-linalool, (E)-ss-farnesene, hexanol, (Z)-3-hexenyl acetate, 4,8-dimethyl-1,3,(E)7-nonatriene, nonanol, the major pheromone component codlemone [(E,E)-8,10-dodecadienol] and the minor pheromone component tetradecanol. Additionally, (E,E)-alpha-farnesene and (Z)-3-hexenol elicited responses specifically in female neurons, whereas (E,E)-farnesol elicited a specific response in a male neuron. Neurons responded to 1-3 odorants, with sometimes overlapping response spectra. A scanning electron microscopic study of the antennae of both sexes supported an earlier study, apart from that long s. trichodea were present in a wreath at the proximal margin of the flagellomere and in addition evenly distributed over the remaining surface, and a previously non-described sensillum type with external basiconic features was revealed, distributed on the proximal and medial region of the flagellomeres.     

Octopamine and tyramine modulate pheromone-sensitive olfactory sensilla of the hawkmoth <Emphasis Type="Italic">Manduca sexta</Emphasis> in a time-dependent manner

In moths octopamine improved pheromone-dependent mate search time dependently. In the nocturnal hawkmoth Manduca sexta long-term tip recordings of trichoid sensilla were performed to investigate whether biogenic amines modulate pheromone transduction time dependently. At three Zeitgebertimes octopamine, tyramine and the octopamine antagonist epinastine were applied during non-adapting pheromone-stimulation. At ZT 8-11, during the photophase, when sensilla were adapted, octopamine and to a lesser extent tyramine increased the bombykal-dependent sensillar potential amplitude and initial action potential (AP) frequency. In addition, during the photophase, when sensilla are less able to resolve pheromone pulses, octopamine rendered pheromone responses more phasic and sensitive, and raised the spontaneous AP frequency. During the late scotophase, at ZT 22-1, when the antenna appeared maximally sensitized for pheromone pulse detection and endogenous octopamine levels are high, exogenously applied octopamine was ineffective. Epinastine blocked the pheromone-dependent AP response at ZT 8-11 and slightly affected it at ZT 22-1, while it had no effect on the sensillar potential amplitude. Epinastine decreased the spontaneous AP activity during photophase and scotophase and rendered pheromone responses more tonic in the scotophase. We hypothesize that the presence of octopamine in the antenna is obligatory for the detection of intermittent pheromone pulses at all Zeitgebertimes.     

No inhibitory effect on receptor neurone activity by sulphur analogues of the sex pheromone component (Z)-5-decenyl acetate in the turnip moth, Agrofis segetum (Lepidoptera: Noctuidae)

Abstract. Olfactory responses from the entire antenna and from single antennal sensilla of the male turnip moth, Agrotis segetum (Lepidoptea: Noctuidae Schiff.), were recorded after stimulation of the antenna with the sex pheromone component, (Z)-5-decenyl acetate (Z5-10:OAc), and three sulphur analogues of this compound. Adaptation of olfactory receptor neurones tuned to Z5-10:OAc was investigated after pre-exposure of these receptor neurones to the key stimulus and to the three sulphur analogues. Both electro-antennographic and single sensillum recordings revealed that the sulphur analogues had a significantly decreased effect compared to the natural stimulus. The pre-exposure experiments demonstrated that no further inhibition of neural activity was observed than could be expected from receptor neurone adaptation. Earlier reports, describing sulphur analogues as possible hyperagonists acting on moth pheromone receptor neurones, are not supported by the present study.     

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。     

Electrophysiological characterisation of olfactory sensilla in the black bean aphid, Aphis fabae

Electrophysiological responses of three different olfactory sensilla (proximal primary rhinaria (PPR), distal primary rhinaria (DPR) and secondary rhinaria (SR)) to the sex pheromone components, nepetalactol, nepetalactone, and a plant volatile, (E)-2-hexenal, were investigated in four different morphs of the black bean aphid, Aphis fabae Scopoli. The DC responses recorded directly from the antennal sensilla and termed electrosensillograms (ESGs) were generally much larger (up to 14 mV) than electroantennogram (EAG) responses (up to 2.5 mV). Characteristic morph-specific response profiles to these compounds were observed in each type of rhinarium and response waveforms were different between (E)-2-hexenal and the sex pheromone components. (E)-2-Hexenal elicited the largest responses at PPR, while nepetalactol and nepetalactone elicited the largest responses at SR in gynoparae and males. Nepetalactol and nepetalactone also showed significant activities on DPR and PPR in all morphs. In contrast, (E)-2-hexenal had almost no activity on SR. However, almost all of the SR investigated in males and gynoparae were sensitive to both nepetalactol and nepetalactone. A small sub-set of male SR responded mainly to nepetalactol. SR of winged virginoparae did not respond to the sex pheromone compounds. Paraffin oil (the solvent control) also elicited significant responses at PPR in virginoparae but not in other morphs. In a further experiment, SR of winged virginoparae showed no response to 30 other plant volatile compounds or the alarm pheromone component, (E)-beta-farnesene. Nepetalactol and nepetalactone had similar dose-response profiles in the SR of gynoparae. The results indicate that SR in males and gynoparae are highly specialised to detect sex pheromone compounds, while the DPR and PPR are relatively broadly tuned to both plant volatiles and sex pheromone components. The presence of SR in winged virginoparae that are not responsive to sex pheromone components, alarm pheromone, or any of the plant volatile compounds tested may indicate a possible role of these sensilla to detect, as yet, unknown compound(s) with a high specificity. The present study also suggests that PPR may play a role in detecting paraffin-related compounds such as cuticular hydrocarbons of plants.