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Pheromones convey rich ethological information and guide insects’ search behavior. Insects navigating in turbulent environments are tasked with the challenge of coding the temporal structure of an odor plume, obliging recognition of the onset and offset of whiffs of odor. The coding mechanisms that shape odor offset recognition remain elusive. We designed a device to deliver sharp pheromone pulses and simultaneously measured the response dynamics from pheromone-tuned olfactory receptor neurons (ORNs) in male moths and Drosophila . We show that concentration-invariant stimulus duration encoding is implemented in moth ORNs by spike frequency adaptation at two time scales. A linear-nonlinear model fully captures the underlying neural computations and offers an insight into their biophysical mechanisms. Drosophila use pheromone cis -vaccenyl acetate (cVA) only for very short distance communication and are not faced with the need to encode the statistics of the cVA plume. Their cVA-sensitive ORNs are indeed unable to encode odor-off events. Expression of moth pheromone receptors in Drosophila cVA-sensitive ORNs indicates that stimulus-offset coding is receptor independent. In moth ORNs, stimulus-offset coding breaks down for short ( < 200 ms) whiffs. This physiological constraint matches the behavioral latency of switching from the upwind surge to crosswind cast flight upon losing contact with the pheromone. Precise odor delivery shows that moth pheromone sensitive olfactory receptor neurons are capable of very fast response to odor offset. Adaptation of the neurons on two different time scales is necessary to reproduce the response time course.

The volatile plant compounds (VPC) alter pheromone perception by insects but mixture effects inside insect olfactory landscapes are poorly understood. We measured the activity of receptor neurons tuned to Z7-12Ac (Z7-ORN), a pheromone component, in the antenna and central neurons in male Agrotis ipsilon while exposed to simple or composite backgrounds of a panel of VPCs representative of the odorant variety encountered by a moth. Maps of activities were built using calcium imaging to visualize which areas in antennal lobes (AL) were affected by VPCs. We compared the VPC activity and their impact as backgrounds at antenna and AL levels, individually or in blends. At periphery, VPCs showed differences in their capacity to elicit Z7-ORN firing response that cannot be explained by differences in stimulus intensities because we adjusted concentrations according to vapor pressures. The AL neuronal network, which reformats the ORN input, did not improve pheromone salience. We postulate that the AL network evolved to increase sensitivity and to encode for fast changes of pheromone at some cost for signal extraction. Comparing blends to single compounds indicated that a blend shows the activity of its most active component. VPC salience seems to be more important than background complexity.

Responses of insect olfactory receptor neurons (ORNs) involve an entry of Ca²⁺ through olfactory heterodimeric receptor complexes. In moths, the termination of ORN responses was found to strongly depend on the external Ca²⁺ concentration through the activation of unknown Ca²⁺-dependent Cl⁻ channels. We thus investigated the molecular identity of these Cl⁻ channels. There is compelling evidence that bestrophins form Cl⁻ channels when expressed in heterologous systems. Here we provide evidence that antennae of the moth Spodoptera littoralis express three transcripts encoding proteins with hallmarks of bestrophins. One of these transcripts, SlitBest1b, is expressed in ORNs. The heterologous expression of SlitBest1b protein in CHO-K1 cells yielded a Ca²⁺-activated Cl⁻ current that shares electrophysiological properties with the native Ca²⁺-activated Cl⁻ current of ORNs. Both currents are anionic, present similar dependence on the intracellular Ca²⁺ concentration, partly inactivate over time, have the same anion permeability sequence, the same sequence of inhibitory efficiency of blockers, the same almost linear I-V relationships and finally both currents do not depend on the cell volume. Therefore, our data suggest that SlitBest1b is a good candidate for being a molecular component of the olfactory Ca²⁺-activated Cl⁻ channel and is likely to constitute part of the insect olfactory transduction pathway. A different function (e.g. regulation of other proteins, maintenance of the anionic homeostasis in the sensillar lymph) and a different role (e.g. involvement in the olfactory system development) cannot be excluded however.

Insect pest management relies mainly on neurotoxic insecticides, including neonicotinoids, leaving residues in the environment. There is now evidence that low doses of insecticides can have positive effects on pest insects by enhancing various life traits. Because pest insects often rely on sex pheromones for reproduction, and olfactory synaptic transmission is cholinergic, neonicotinoid residues could modify chemical communication. We recently showed that treatments with different sublethal doses of clothianidin could either enhance or decrease behavioural sex pheromone responses in the male moth, Agrotis ipsilon. We investigated now effects of the behaviourally active clothianidin doses on the sensitivity of the peripheral and central olfactory system. We show with extracellular recordings that both tested clothianidin doses do not influence pheromone responses in olfactory receptor neurons. Similarly, in vivo optical imaging does not reveal any changes in glomerular response intensities to the sex pheromone after clothianidin treatments. The sensitivity of intracellularly recorded antennal lobe output neurons, however, is upregulated by a lethal dose 20 times and downregulated by a dose 10 times lower than the lethal dose 0. This correlates with the changes of behavioural responses after clothianidin treatment and suggests the antennal lobe as neural substrate involved in clothianidin-induced behavioural changes.

Responses of insect olfactory receptor neurons (ORNs) involve an entry of Ca.sup.2+ through olfactory heterodimeric receptor complexes. In moths, the termination of ORN responses was found to strongly depend on the external Ca.sup.2+ concentration through the activation of unknown Ca.sup.2+ -dependent Cl.sup.- channels. We thus investigated the molecular identity of these Cl.sup.- channels. There is compelling evidence that bestrophins form Cl.sup.- channels when expressed in heterologous systems. Here we provide evidence that antennae of the moth Spodoptera littoralis express three transcripts encoding proteins with hallmarks of bestrophins. One of these transcripts, SlitBest1b, is expressed in ORNs. The heterologous expression of SlitBest1b protein in CHO-K1 cells yielded a Ca.sup.2+ -activated Cl.sup.- current that shares electrophysiological properties with the native Ca.sup.2+ -activated Cl.sup.- current of ORNs. Both currents are anionic, present similar dependence on the intracellular Ca.sup.2+ concentration, partly inactivate over time, have the same anion permeability sequence, the same sequence of inhibitory efficiency of blockers, the same almost linear I-V relationships and finally both currents do not depend on the cell volume. Therefore, our data suggest that SlitBest1b is a good candidate for being a molecular component of the olfactory Ca.sup.2+ -activated Cl.sup.- channel and is likely to constitute part of the insect olfactory transduction pathway. A different function (e.g. regulation of other proteins, maintenance of the anionic homeostasis in the sensillar lymph) and a different role (e.g. involvement in the olfactory system development) cannot be excluded however.

The response of insect olfactory receptor neurons (ORN) involves an increase in intracellular Ca^sup 2+^ concentration, as in vertebrate ORNs. In order to decipher the Ca^sup 2+^ clearance mechanisms in insect ORNs, we have investigated the presence of a plasma membrane Ca^sup 2+^ ATPase (PMCA) in the peripheral olfactory system of the moth Spodoptera littoralis. From an analysis of a male antennal expressed-sequence-tag database combined with a strategy of 5'/3' rapid amplification of cDNA ends plus the polymerase chain reaction, we have cloned a full-length cDNA encoding a PMCA. In adult males, the PMCA transcript has been found in various tissues, including the antennae in which its presence has been detected in the sensilla trichodea, and in cultured ORNs. The PMCA gene is slightly expressed at the end of the pupal stage, reaches a maximum at emergence and is maintained at a high level during the adult period. Taken together, these results provide, for the first time, molecular evidence for the putative participation of a PMCA in signalling pathways responsible for the establishment and functioning of the insect peripheral olfactory system. [PUBLICATION ABSTRACT]

The response of insect olfactory receptor neurons (ORNs) involves an increase in intracellular [Ca.sup.2+] concentration, as in vertebrate ORNs. In order to decipher the [Ca.sup.2+] clearance mechanisms in insect ORNs, we have investigated the presence of a plasma membrane [Ca.sup.2+] ATPase (PMCA) in the peripheral olfactory system of the moth Spodoptera littoralis. From an analysis of a male antennal expressed-sequence-tag database combined with a strategy of 5'/3' rapid amplification of cDNA ends plus the polymerase chain reaction, we have cloned a full-length cDNA encoding a PMCA. In adult males, the PMCA transcript has been found in various tissues, including the antennae in which its presence has been detected in the sensilla trichodea, and in cultured ORNs. The PMCA gene is slightly expressed at the end of the pupal stage, reaches a maximum at emergence and is maintained at a high level during the adult period. Taken together, these results provide, for the first time, molecular evidence for the putative participation of a PMCA in signalling pathways responsible for the establishment and functioning of the insect peripheral olfactory system. Keywords Plasma membrane calcium ATPase Olfactory transduction * Calcium clearance * Olfactoryreceptor neuron * Spodoptera littoralis (Insecta)

The response of insect olfactory receptor neurons (ORNs) involves an increase in intracellular Ca(2+) concentration, as in vertebrate ORNs. In order to decipher the Ca(2+) clearance mechanisms in insect ORNs, we have investigated the presence of a plasma membrane Ca(2+) ATPase (PMCA) in the peripheral olfactory system of the moth Spodoptera littoralis. From an analysis of a male antennal expressed-sequence-tag database combined with a strategy of 5'/3' rapid amplification of cDNA ends plus the polymerase chain reaction, we have cloned a full-length cDNA encoding a PMCA. In adult males, the PMCA transcript has been found in various tissues, including the antennae in which its presence has been detected in the sensilla trichodea, and in cultured ORNs. The PMCA gene is slightly expressed at the end of the pupal stage, reaches a maximum at emergence and is maintained at a high level during the adult period. Taken together, these results provide, for the first time, molecular evidence for the putative participation of a PMCA in signalling pathways responsible for the establishment and functioning of the insect peripheral olfactory system.

The response of insect olfactory receptor neurons (ORNs) involves an increase in intracellular Ca super(2+) concentration, as in vertebrate ORNs. In order to decipher the Ca super(2+) clearance mechanisms in insect ORNs, we have investigated the presence of a plasma membrane Ca super(2+) ATPase (PMCA) in the peripheral olfactory system of the moth Spodoptera littoralis. From an analysis of a male antennal expressed-sequence-tag database combined with a strategy of 5'/3' rapid amplification of cDNA ends plus the polymerase chain reaction, we have cloned a full-length cDNA encoding a PMCA. In adult males, the PMCA transcript has been found in various tissues, including the antennae in which its presence has been detected in the sensilla trichodea, and in cultured ORNs. The PMCA gene is slightly expressed at the end of the pupal stage, reaches a maximum at emergence and is maintained at a high level during the adult period. Taken together, these results provide, for the first time, molecular evidence for the putative participation of a PMCA in signalling pathways responsible for the establishment and functioning of the insect peripheral olfactory system.

The response of insect olfactory receptor neurons (ORNs) involves an increase in intracellular Ca2+ concentration, as in vertebrate ORNs. In order to decipher the Ca2+ clearance mechanisms in insect ORNs, we have investigated the presence of a plasma membrane Ca2+ ATPase (PMCA) in the peripheral olfactory system of the moth Spodoptera littoralis. From an analysis of a male antennal expressed-sequence-tag database combined with a strategy of 5'/3' rapid amplification of cDNA ends plus the polymerase chain reaction, we have cloned a full-length cDNA encoding a PMCA. In adult males, the PMCA transcript has been found in various tissues, including the antennae in which its presence has been detected in the sensilla trichodea, and in cultured ORNs. The PMCA gene is slightly expressed at the end of the pupal stage, reaches a maximum at emergence and is maintained at a high level during the adult period. Taken together, these results provide, for the first time, molecular evidence for the putative participation of a PMCA in signalling pathways responsible for the establishment and functioning of the insect peripheral olfactory system.