Dancing in the purple rain: color affinity and oviposition choices in Aedes sierrensis (Diptera: Culicidae).

The western tree hole mosquito, Aedes sierrensis (Ludlow) (Diptera: Clucidae), is a pestiferous mosquito with a range extending over the entire pacific seaboard and into portions of the intermountain west. As a peridomestic heartworm vector, it demands at least some level of surveillance to understand its abundance. However, the species is refractory to a majority of conventional vector surveillance approaches for tracking mosquitoes. To find more options for Aedes sierrensis surveillance, a variety of oviposition attractants were evaluated in arena-style choice assays using colony reared adults. A range of infusion treatments (e.g., alfalfa, oak, and beetroot) were examined and then combined with investigations of liquid color as well as ovicup color and entryway position. These studies revealed that Ae. sierrensis have an affinity for purple coloration, plain water, and larger entryway sizes for oviposition cups. A prototype ovicup was 3D-printed using purple filament and multiple types of entryways, and used to re-test infusion waters. No particular attraction differences were detected after normalizing for purple color. Comparisons to black 3D-printed cups yielded surprising observations that male mosquitoes also aggregated on purple cups while females sheltered, but not necessarily oviposited, in black cups. Although this was only a laboratory-based assessment, these studies provide useful information for future field trials of potential oviposition traps for surveillance of Ae. sierrensis.

Illness and femininity in Hilary Mantel's Giving Up the Ghost (2003).

This paper offers a reading of Hilary Mantel's memoir, Giving Up The Ghost (2003). The interest of the memoir derives from the fact that it provides an exceptionally rich picture of the impact of family life on a child's attitudes towards her own body. Mantel presents her bodily experiences as primitive, often unconscious, perceptions of the relationships within her family of origin. When she discovers new things about those relationships, she must register the change through her body in some way. Drawing on a range of concepts taken from psychoanalytic psychosomatics, I suggest that at the heart of the memoir is the author's bafflement at the repeated and uncanny irruption of a conflict between her body as a somewhat autonomous signifying entity and the psychological strength she seeks and often finds through identifications with family members. I argue that this conflict overlapped with her acceptance of a female gender identity. The sustained nature of this conflict prevented her from establishing a metric of what I will call 'psychosomatic normality', with disastrous consequences when she began to suffer the symptoms of acute endometriosis. The memoir also shows the power of early life in determining how diseases are experienced subjectively, over time.

Moth pheromone-selective projection neurons with cell bodies in the antennal lobe lateral cluster exhibit diverse morphological and neurophysiological characteristics.

Olfactory projection neurons convey information from the insect antennal lobe (AL) to higher brain centers. Previous reports have demonstrated that pheromone-responsive projection neurons with cell bodies in the moth medial cell cluster (mcPNs) predominantly have dendritic arborizations in the sexually dimorphic macroglomerular complex (MGC) and send an axon from the AL to the calyces of the mushroom body (CA) as well as the lateral horn (LH) of the protocerebrum via the medial AL tract. These neurons typically exhibit a narrow odor tuning range related to the restriction of their dendritic arbors within a single glomerulus (uniglomerular). In this study, we report on the diverse physiological and morphological properties of a group of pheromone-responsive olfactory projection neurons with cell bodies in the AL lateral cell cluster (MGC lcPNs) of two closely related moth species. All pheromone-responsive lcPNs appeared to exhibit "basket-like" dendritic arborizations in two MGC compartments and made connections with various protocerebral targets including ventrolateral and superior neuropils via projections primarily through the lateral AL tract and to a lesser extent the mediolateral antennal lobe tract. Physiological characterization of MGC lcPNs also revealed a diversity of response profiles including those either enhanced by or reliant upon presentation of a pheromone blend. These responses manifested themselves as higher maximum firing rates and/or improved temporal resolution of pulsatile stimuli. MGC lcPNs therefore participate in conveying diverse olfactory information relating to qualitative and temporal facets of the pheromone stimulus to a more expansive number of protocerebral targets than their mcPN counterparts.

Animal Communication: When I'm Calling You, Will You Answer Too?

Male moths compete to arrive first at a female releasing pheromone. A new study reveals that additional pheromone cues released only by younger females may prompt males to avoid them in favor of older but more fecund females.

The body in Martin Amis's Experience (2000).

This paper focuses on the presentation of the body in Martin Amis's memoir Experience (2000) and compares Amis's account of the growth of his mind and body with ideas put forward by writers in the phenomenological and psychoanalytic traditions. Using the ideas of body schema, projective identification and idea of safety, it advances a new conceptual framework for the thinking about the contribution that the body makes to selfhood in autobiography.

Transplant Antennae and Host Brain Interact to Shape Odor Perceptual Space in Male Moths.

Behavioral responses to odors rely first upon their accurate detection by peripheral sensory organs followed by subsequent processing within the brain's olfactory system and higher centers. These processes allow the animal to form a unified impression of the odor environment and recognize combinations of odorants as single entities. To investigate how interactions between peripheral and central olfactory pathways shape odor perception, we transplanted antennal imaginal discs between larval males of two species of moth Heliothis virescens and Heliothis subflexa that utilize distinct pheromone blends. During metamorphic development olfactory receptor neurons originating from transplanted discs formed connections with host brain neurons within olfactory glomeruli of the adult antennal lobe. The normal antennal receptor repertoire exhibited by males of each species reflects the differences in the pheromone blends that these species employ. Behavioral assays of adult transplant males revealed high response levels to two odor blends that were dissimilar from those that attract normal males of either species. Neurophysiological analyses of peripheral receptor neurons and central olfactory neurons revealed that these behavioral responses were a result of: 1. the specificity of H. virescens donor olfactory receptor neurons for odorants unique to the donor pheromone blend and, 2. central odor recognition by the H. subflexa host brain, which typically requires peripheral receptor input across 3 distinct odor channels in order to elicit behavioral responses.

Opium as a Literary Stimulant: The Case of Samuel Taylor Coleridge.

In our era, the idea of a stimulant is synonymous with its biochemical properties. A stimulant, we think, is a substance that enhances the activity of the central and peripheral nervous systems. But in the eighteenth century, a new family of theories about the workings of stimulants took shape, based on exciting but erroneous assumptions. Proponents of these theories thought that many more diseases were "nervous" in origin than had previously been supposed. They hoped that the workings of the "nervous power" could be aided by the judicious use of stimulants and narcotics. Practitioners working within this broad "neuropathological" paradigm equated the workings of stimulation with those of gravity. Stimulation, they believed, was a kind of master principle in nature. Some hoped it would help refound medicine on Newtonian, mathematical lines. For patients, the most visible legacy of the neuropathological revolution was the abandonment of bloodletting or "cupping" and the increasingly widespread use of opium and alcohol in medical treatments. In this chapter, I explore the career of one of the most famous writers of the Romantic era, Samuel Taylor Coleridge (1772-1834) who had the misfortune to live through this therapeutic revolution. I describe the circumstances under which he came to take opiates and the development of his opinions about their effect on him.

Male moths optimally balance take-off thoracic temperature and warm-up duration to reach a pheromone source quickly.

Animal activities, such as foraging and reproduction, are constrained by decisions about how to allocate energy and time efficiently. Overall, male moths invest less in reproduction than females, but they are thought to engage in a scramble competition for access to females that advertise readiness to mate by releasing sexual pheromones. However, before male moths can follow the pheromone, they often need to heat their flight muscles by shivering to produce sufficient power for sustained flight. Here, we show that Helicoverpa zea males that sense the female pheromone at high ambient temperatures take off with higher thoracic temperature, shiver for less time and warm up faster than males tested at lower ambient temperatures. These higher take-off temperatures translate into higher airspeeds, underscoring the importance of thoracic temperature for flight performance. Furthermore, shorter combined duration for warm-up and pheromone-mediated optomotor anemotaxis is consistent with the idea that males engage in scramble competition for access to females in nature. Our results strongly suggest that male moths minimize the time between perceiving the female's pheromone signal and arriving at the source by optimizing thermoregulatory behaviour and temperature-dependent flight performance in accordance with ambient temperature conditions. Our finding that moths engage in a trade-off between rapid flight initiation and suboptimal flight performance suggests a sensorimotor control mechanism that involves a complex interaction with the thermal environment.

Female pheromones modulate flight muscle activation patterns during preflight warm-up.

At low ambient temperature Helicoverpa zea male moths engage in warm-up behavior prior to taking flight in response to an attractive female pheromone blend. Male H. zea warm up at a faster rate when sensing the attractive pheromone blend compared with unattractive blends or blank controls (Crespo et al. 2012), but the mechanisms involved in this olfactory modulation of the heating rate during preflight warm-up are unknown. Here, we test three possible mechanisms for increasing heat production: 1) increased rate of muscle contraction; 2) reduction in mechanical movement by increased overlap in activation of the antagonistic flight muscles; and 3) increased activation of motor units. To test which mechanisms play a role, we simultaneously recorded electrical activation patterns of the main flight muscles (dorsolongitudinal and dorsoventral muscles), wing movement, and thoracic temperature in moths exposed to both the attractive pheromone blend and a blank control. Results indicate that the main mechanism responsible for the observed increase in thoracic heating rate with pheromone stimulation is the differential activation of motor units during each muscle contraction cycle in both antagonistic flight muscles. This additional activation lengthens the contracted state within each cycle and thus accounts for the greater heat production. Interestingly, the rate of activation (frequency of contraction cycles) of motor units, which is temperature dependent, did not vary between treatments. This result suggests that the activation rate is determined by a temperature-dependent oscillator, which is not affected by the olfactory stimulus, but activation of motor units is modulated during each cycle.

Pheromone mediated modulation of pre-flight warm-up behavior in male moths.

An essential part of sexual reproduction typically involves the identification of an appropriate mating partner. Males of many moth species utilize the scent of sex pheromones to track and locate conspecific females. However, before males engage in flight, warm-up by shivering of the major flight muscles is necessary to reach a thoracic temperature suitable to sustain flight. Here we show that Helicoverpa zea males exposed to an attractive pheromone blend (and in some instances to the primary pheromone component alone) started shivering earlier and took off at a lower thoracic temperature than moths subjected to other incomplete or unattractive blends. This resulted in less time spent shivering and faster heating rates. Two interesting results emerge from these experiments. First, the rate of heat generation can be modulated by different olfactory cues. Second, males detecting the pheromone blend take off at lower thoracic temperatures than males exposed to other stimuli. The take-off temperature of these males was below that for optimal power production in the flight muscles, thus generating a trade-off between rapid departure and suboptimal flight performance. Our results shed light on thermoregulatory behaviour of unrestrained moths associated with the scramble competition for access to females and suggest ecological trade-offs between rapid flight initiation and sub-optimal flight performance.

Antennal lobe organization in the slender pigeon louse, Columbicola columbae (Phthiraptera: Ischnocera).

This study reports on the structure of the antennal lobe of the pigeon louse, Columbicola columbae. Anterograde staining of antennal receptor neurons revealed an antennal lobe with a few diffuse compartments, an organization distinct from the typical spheroidal glomerular structure found in the olfactory bulb of vertebrates and the antennal lobe of many other insects. This anatomical arrangement of neuronal input is somewhat reminiscent of the aglomerular antennal lobe previously reported in psyllids and aphids. As in psyllids, reports on the odor-mediated behavior of C. columbae suggest that the olfactory sense is important in these animals and indicates that a glomerular organization of the antennal lobe may not be necessary to subtend odor-mediated behaviors in all insects. The diffuse or aglomerular antennal lobe organization found in these two Paraneopteran insect orders might represent an independently evolved reduction due to similar ecological constraints.

Hairpencil volatiles influence interspecific courtship and mating between two related moth species.

Reproductive isolation between sympatric, closely related species can be accomplished through a variety of pre-zygotic isolating mechanisms, including courtship-signaling behavior that involves pheromones. In the moths Heliothis virescens and H. subflexa, males display abdominal hairpencils (HP), which release volatile chemicals during courtship. In this study, we demonstrated that HP volatiles released by male H. subflexa function to improve mating success with conspecific females. Interspecific mating experiments were conducted to determine any influence of HP volatiles on species isolation. Female H. virescens and H. subflexa were observed during courtship with males of the other species, following either sham-operation or ablation of HPs, both with and without concurrent presentation of HP volatiles. Mating success was improved by co-presentation of HP extract from males of the same species during courtship. Ablation of HPs improved mating between H. subflexa females and H. virescens males. During interspecific matings, male H. virescens attempted copulation less frequently in the presence of H. virescens HP extract, though H. subflexa males were not affected by the presence of H. subflexa HP extract. This suggests that HP volatiles produced by males of each species may inhibit mating between species through effects on males (H. virescens) and females (H. subflexa).

Sexual isolation of male moths explained by a single pheromone response QTL containing four receptor genes.

Long distance sexual communication in moths has fascinated biologists because of the complex, precise female pheromone signals and the extreme sensitivity of males to specific pheromone molecules. Progress has been made in identifying some genes involved in female pheromone production and in male response. However, we have lacked information on the genetic changes involved in evolutionary diversification of these mate-finding mechanisms that is critical to understanding speciation in moths and other taxa. We used a combined quantitative trait locus (QTL) and candidate gene approach to determine the genetic architecture of sexual isolation in males of two congeneric moths, Heliothis subflexa and Heliothis virescens. We report behavioral and neurophysiological evidence that differential male responses to three female-produced chemicals (Z9-14:Ald, Z9-16:Ald, Z11-16:OAc) that maintain sexual isolation of these species are all controlled by a single QTL containing at least four odorant receptor genes. It is not surprising that pheromone receptor differences could control H. subflexa and H. virescens responses to Z9-16:Ald and Z9-14:Ald, respectively. However, central rather than peripheral level control over the positive and negative responses of H. subflexa and H. virescens to Z11-16:OAc had been expected. Tight linkage of these receptor genes indicates that mutations altering male response to complex blends could be maintained in linkage disequilibrium and could affect the speciation process. Other candidate genes such as those coding for pheromone binding proteins did not map to this QTL, but there was some genetic evidence of a QTL for response to Z11-16:OH associated with a sensory neuron membrane protein gene.

Thomas Beddoes and the German psychological tradition.

This paper considers Thomas Beddoes's role in disseminating German psychological ideas in Britain. It describes the German tradition as inaugurated by Karl Philipp Moritz (1756-93) and considers the chief differences between this tradition and the English one stemming from David Hartley. It is suggested that Beddoes found strong support for his convictions about human interiority in writings by Moritz and his followers. In particular, these enabled him to think about sanity and madness as being continuous with one another (rather than as one another's negation); they helped him locate the signs of madness in ordinary childhood behaviours; they reinforced his suspicions that many so-called nervous disorders were psychically caused; and they supplied him with a conception of unconscious passion. The paper concludes by considering Beddoes's appeal to Shakespeare's plays as a source of clinical knowledge about the nature of insanity, and argues that Beddoes has been overlooked as a crucial source for nineteenth-century psychiatrist-bardologists such as J. Conolly, J. C. Bucknill and H. Maudsley.

Phenology of semiochemical-mediated host foraging by the western boxelder bug, Boisea rubrolineata, an aposematic seed predator.

The western boxelder bug (BEB), Boisea rubrolineata (Heteroptera: Rhopalidae), is a specialist herbivore of boxelder trees, Acer negundo. We tested the hypothesis that BEBs use semiochemicals to locate host trees. Headspace volatiles from trees bearing staminate inflorescences ("staminate trees") and from trees bearing pistillate inflorescences ("pistillate trees") were collected throughout the season and bioassayed in Y-tube olfactometer experiments. Headspace extracts of early-season, pollen-bearing staminate trees and midseason pistillate trees with mature samaras (seed pods) attracted female and male BEBs. By using coupled gas chromatographic-electroantennographic detection and gas chromatography-mass spectrometry, we identified and tested a five-component synthetic blend of candidate semiochemicals (hexanol, pentyl acetate, phenylacetonitrile, 2-phenethyl acetate, and trans-nerolidol). This blend attracted females, males, and fifth-instar nymphs. Phenylacetonitrile by itself was as attractive as the five-component blend to both adults and nymphs. By responding to phenylacetonitrile emitted by pollen-bearing staminate trees and pistillate trees with maturing seeds, BEBs appear to track and exploit the availability of nutrient-rich food sources, suggesting that the bugs' reproductive ecology is synchronized to the phenology of their host boxelder tree.

Central processing of natural odor mixtures in insects.

In nature, virtually all olfactory stimuli are mixtures of many single odorants. Behavioral experiments repeatedly have demonstrated that an animal's olfactory system is capable of discriminating behaviorally relevant from irrelevant odor mixtures. However, the sensory mechanisms that underlie such discriminative capability remain elusive. The limited anatomical and physiological evidence collected from both insect and vertebrate models that pertains to this topic is scattered in the literature dating back to early 1980s. Thus, a synthesis of this information that includes more recent findings is needed in order to provide a basis for probing the fundamental question from a new angle. In this review, we discuss several proposed models for mixture processing, along with experimental data gathered from both the initial stage of olfactory processing (i.e., antennal lobe in insects or olfactory bulb in vertebrates) and higher areas of the brain, with an emphasis on how the lateral circuits in the antennal lobe or olfactory bulb may contribute to mixture processing. Based on empirical data as well as theoretical modeling, we conclude that odor mixtures may be represented both at the single-neuron level and at the population level. The difference between these two types of processing may reside in the degree of plasticity, with the former being hard-wired and the latter being more subjected to network modulation.

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.

Inheritance of olfactory preferences I. Pheromone-mediated behavioral responses of Heliothis subflexa x Heliothis virescens hybrid male moths.

Shifts in male preference for qualitatively different pheromone blends appear to have played a fundamental role in the divergence of olfactory communication and evolution of moth species. As an initial step in documenting the genetic complexity underlying such shifts, we characterized the behavioral responses of hybrid male moths created by mating two heliothine moth species, Heliothis subflexa and Heliothis virescens. Between 67 and 96% of hybrid males flew upwind and contacted the pheromone source when presented with a blend consisting of (Z)-11-hexadecenal (Z11-16:Ald), (Z)-9-hexadecenal (Z9-16:Ald), and (Z)-11-hexadecenol (Z11-16:OH) in a 1:0.5:0.1 ratio that has previously been shown to be attractive to H. subflexa males. In addition, an H. virescens blend of Z11-16:Ald and (Z)-9-tetradecenal (Z9-14:Ald) enhanced by the addition of Z11-16:OH (in a 1:0.05:0.1 mixture) was attractive to hybrid males (26-64% source contact), but significantly fewer males reached the odor source compared to the blend containing Z9-16:Ald. A blend in which the dosage of Z9-14:Ald was doubled, however, was equally attractive (75-77% source contact) as the Z9-16:Ald-containing blend. Consecutive presentation of two blends revealed that individual hybrid males responded equally well to blends containing either Z9-14:Ald or Z9-16:Ald. Together these results suggest that in addition to Z11-16:Ald, hybrid males: (1) required either Z9-16:Ald (likeH. subflexamales) or Z9-14:Ald (like H. virescens males); (2) required the presence of Z11-16:OH (H. subflexa dominant); (3) were not adversely affected by the presence of Z11-16:Ac (H. subflexa dominant). The behavioral response phenotype of hybrid males was therefore influenced by genetic factors inherited from both parental species.