Thomas C. Baker

Odor Detection in Insects: Volatile Codes

Insect olfactory systems present models to study interactions between animal genomes and the environment. They have evolved for fast processing of specific odorant blends and for general chemical monitoring. Here, we review molecular and physiological mechanisms in the context of the ecology of chemical signals. Different classes of olfactory receptor neurons (ORNs) detect volatile chemicals with various degrees of specialization. Their sensitivities are determined by an insect-specific family of receptor genes along with other accessory proteins. Whereas moth pheromones are detected by highly specialized neurons, many insects share sensitivities to chemical signals from microbial processes and plant secondary metabolism. We promote a more integrated research approach that links molecular physiology of receptor neurons to the ecology of odorants.

Sexual Communication with Pheromones

A critical event in sexual reproduction is location or recruitment of a mate. In a number of insect groups, the necessary movements in time and space are often mediated by pheromones. One sex may recruit the other, or both sexes may be attracted to the chemical emitters. Aggregation may be viewed as the end result of movement reactions that reduce the distance between individuals in their environment. Such clustering may be brought about by a combination of attraction and arrestment, which are themselves not orientation mechanisms but rather end results, i.e., displacements, caused by movement reactions (Kennedy, 1978). For sex- and aggregation-pheromone communication, we define attraction as the net displacement of one individual toward the chemical source. Conversely, arrestment is the lack of net displacement toward or away from the chemical source. Both displacement phenomena may be viewed as part of a continuum caused by pheromone mediation of quite disparate movement reactions, such as orthokinesis, klinotaxis and anemotaxis (see Bell, Chapter 4 and Carde, Chapter 5). That attraction and arrestment are only outcomes, not mechanisms, does not diminish the heuristic value of these terms; they are a capsule summary of the change in spacing between an individual and the chemical source. To an organism responding to sex pheromone, proximate cues and orientation mechanisms notwithstanding, such outcomes are the result of evolutionary selection.

Odour-plume dynamics influence the brain's olfactory code

The neural computations used to represent olfactory information in the brain have long been investigated. Recent studies in the insect antennal lobe suggest that precise temporal and/or spatial patterns of activity underlie the recognition and discrimination of different odours, and that these patterns may be strengthened by associative learning. It remains unknown, however, whether these activity patterns persist when odour intensity varies rapidly and unpredictably, as often occurs in nature. Here we show that with naturally intermittent odour stimulation, spike patterns recorded from moth antennal-lobe output neurons varied predictably with the fine-scale temporal dynamics and intensity of the odour. These data support the hypothesis that olfactory circuits compensate for contextual variations in the stimulus pattern with high temporal precision. The timing of output neuron activity is constantly modulated to reflect ongoing changes in stimulus intensity and dynamics that occur on a millisecond timescale.

Effects of intermittent and continuous pheromone stimulation on the flight behaviour of the oriental fruit moth, Grapholita molesta

ABSTRACT. When male oriental fruit moths, Grapholita molesta (Busck) (Tortricidae), casting in clean air entered an airstream permeated with pheromone their flight tracks changed immediately on initial contact with pheromone, but after a few seconds returned to casting as if in clean air. The degree of change in the flight track was directly related to the concentration of pheromone. Although little net uptunnel movement occurred in response to the continuous stimulation provided by a uniformly permeated airstream, when an intermittent stimulus provided by a point‐source plume was superimposed onto the permeated airstream moths were able to ‘lock on’ and zigzag uptunnel in the plume. The percentage of moths doing so corresponded to the difference between the peak concentration within the plume and the background concentration of pheromone permeating the airstream. Moths also locked onto, and flew upwind along the pheromone‐clean‐air boundary formed along a pheromone‐permeated side corridor. Because a similar response was observed along a horizontal edge between a pheromone‐permeated floor corridor and clean air, we conclude that the intermittent stimulation at the edge perpetuated the narrow zigzagging response to pheromone.

Host plant volatiles synergize responses of sex pheromone-specific olfactory receptor neurons in male Helicoverpa zea

Abstract. 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 neurons 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.

Olfactory reactions of the twelve-spotted lady beetle, Coleomegilla maculata and the green lacewing, Chrysoperla carnea to semiochemicals released from their prey and host plant : Electroantennogram and behavioral responses

Electroantennograms (EAGs) were recorded from two predatory insect species, the twelve-spotted lady beetle, Coleomegilla maculata and the green lacewing, Chrysoperla carnea in response to semiochemicals emitted from one of their prey species, the pea aphid Acyrthosiphon pisum and their host plant. EAG responses were also recorded from C. maculata in response to extracts from individuals of the opposite sex and to extracts from an herbaceous plant, catnip Nepeta cataria. Extracts of catnip and two sex pheromone components of aphids, (4aS,7S,7aR)-nepetalactone and (1R,4aS,7S,7aR)-nepetalactol, elicited significant EAG responses from the antennae of both predatory species. Of 10 corn volatile compounds tested, C. carnea adults responded most strongly to 2-phenylethanol and (E)-β-farnesene. A significant difference in EAG response to extracts of corn leaf collections was observed between male and female C. carnea. In C. maculata, significant EAG responses were elicited by most of the tested corn volatile compounds, except α-pinene and (E)-2-hexenal. The highest EAG responses were observed in response to (E)-β-farnesene, α-terpineol, 2-phenylethanol, and β-caryophyllene. Sexual differences in EAG responses of C. maculata were only found in response to 1-octen-3-ol. Male antennae of C. maculata produced significant EAG responses to extracts from conspecific females, but not to males, which indicates that some chemicals from females could be involved in sexual communication. A significant EAG response also was recorded in response to the extracts of fluids produced during “reflex bleeding.” Male and female antennae of both species exhibited similar dose–response curves to most of the selected compounds, although female C. maculata antennae exhibited higher thresholds in response to several compounds including α-terpineol, (Z)-3-hexenol, and (4aS,7S,7aR)-nepetalactone. Field tests showed that 2-phenylethanol was highly attractive to both sexes of the two investigated species. Only C. maculata was attracted to traps baited with α-terpineol.

Two sex pheromone components of the tobacco budworm moth, Heliothisvirescens

Abstract Two compounds were isolated from female Heliothis virescens (Lepidoptera: Noctuidae) extracts and identified as cis -9-tetradecenal and cis -ll-hexadecenal. Together they elicit intense male H . virescnes response in laboratory tests and have attracted males in the field. Although cis -ll-hexadecenal is an H . zea sex pheromone, no evidence was obtained for cis -9-tetradecenal in H . zea .

A pulsed cloud of sex pheromone elicits upwind flight in male moths

ABSTRACT. Male oriental fruit moths do not fly upwind in a continuous uniform cloud of pheromone, but readily do so when the cloud is pulsed at 1 or 0.5/s or when a plume from a point source of pheromone is placed within the continuous cloud. It is suggested that males of moth species that require such fluctuating pheromone stimulation for upwind flight will normally receive it from a filamentous, point‐source‐produced plume. However, we hypothesize that upwind progress may cease close to the source due to excessively high emission rates or inappropriate blend ratios, when fluctuating sensory output becomes attenuated, despite higher actual molecular concentration fluctuations.

Endogenous and exogenous factors affecting periodicities of female calling and male sex pheromone response in Grapholitha molesta (Busck)

In Grapholitha molesta periodicities of both female calling and male response to sex pheromone by wing fanning while walking were determined in part by circadian rhythms. The lights-on photoperiodic cue was at least partly responsible for setting the phase of the female calling rhythm. Absolute temperature levels and not necessarily a decrease in temperature modified the timing of calling; there were both high and low thresholds of temperature and, at particular photoperiod times, a temperature range optimal for calling. When the previous performance of calling was prevented by subthreshold temperatures, calling during the next period commenced earlier if temperatures were favourable. Thus, the previous performance of calling may establish a refractory period, or temperature decrease may act as a cue resetting the phase of the calling rhythm. The ability to use both an endogenous clock and exogenous temperature cues to synchronize sexual activity appears adaptive for a temperate zone insect whose multiple generations are exposed to both long periods of favourable climatic conditions in summer and harsh, unpredictable conditions in spring or fall.

Functional morphology of antennal chemoreceptors of the parasitoid Microplitis croceipes (Hymenoptera: Braconidae)

The specialist parasitoid, Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), uses chemical cues from plants damaged by herbivore-feeding and also plant by-products in host location and acceptance. These chemicals are detected by the wasps antennae. We conducted scanning and transmission electron microscopy studies of male and female M. croceipes antennae and detected no distinct morphological differences in the chemoreceptors between the sexes. Male antennae are approximately twice as long as female antennae. We found five morphological sensillar types in both sexes: sensilla (s) trichodea were the most abundant and distributed over the whole antenna; s. chaetica were present in low numbers only on the scape and pedicel; and s. basiconica, s. coeloconica and elongated s. placodea were found only on the flagellum. Ultrastructural investigations revealed pore systems on s. basiconica and s. placodea. In s. placodea, sensory neurons run parallel to the longitudinal axis of the sensilla with terminal pores. We recorded responses from single olfactory receptor neurons in s. placodea to two plant-emitted volatiles, cis-3-hexenol and ocimene, and two anthropogenic compounds, cyclohexanone and 2-diisopropylaminoethanol. Male receptor neurons were more sensitive than those of females with significantly higher spike frequency being registered from neurons in males.