Chemical Purification of Terbium-155 from Pseudo-Isobaric Impurities in a Mass Separated Source Produced at CERN.

Four terbium radioisotopes (149, 152, 155, 161Tb) constitute a potential theranostic quartet for cancer treatment but require any derived radiopharmaceutical to be essentially free of impurities. Terbium-155 prepared by proton irradiation and on-line mass separation at the CERN-ISOLDE and CERN-MEDICIS facilities contains radioactive 139Ce16O and also zinc or gold, depending on the catcher foil used. A method using ion-exchange and extraction chromatography resins in two column separation steps has been developed to isolate 155Tb with a chemical yield of ≥95% and radionuclidic purity ≥99.9%. Conversion of terbium into a form suitable for chelation to targeting molecules in diagnostic nuclear medicine is presented. The resulting 155Tb preparations are suitable for the determination of absolute activity, SPECT phantom imaging studies and pre-clinical trials.

Bed bug aggregation on dirty laundry: a mechanism for passive dispersal.

Bed bugs have shown a recent and rapid global expansion that has been suggested to be caused by cheap air travel. How a small, flightless and anachoretic insect that hides within its host's sleeping area manages to travel long distances is not yet clear. Bed bugs are attracted to the odour of sleeping humans and we suggest that soiled clothing may present a similarly attractive cue, allowing bed bugs to 'hitch-hike' around the world after aggregating in the laundry bags of travellers. We show that (1) soiled clothing is significantly more attractive than clean clothing to active bed bugs moving within a bedroom sized arena and (2) elevation of CO2 to a level that simulates human occupancy in the same arena appears to initiate search behaviour rather than direct it. Our results show, for the first time, how leaving worn clothing exposed in sleeping areas when travelling can be exploited by bed bugs to facilitate passive dispersal.

Use of habitat odour by host-seeking insects.

Locating suitable feeding or oviposition sites is essential for insect survival. Understanding how insects achieve this is crucial, not only for understanding the ecology and evolution of insect-host interactions, but also for the development of sustainable pest-control strategies that exploit insects' host-seeking behaviours. Volatile chemical cues are used by foraging insects to locate and recognise potential hosts but in nature these resources usually are patchily distributed, making chance encounters with host odour plumes rare over distances greater than tens of metres. The majority of studies on insect host-seeking have focussed on short-range orientation to easily detectable cues and it is only recently that we have begun to understand how insects overcome this challenge. Recent advances show that insects from a wide range of feeding guilds make use of 'habitat cues', volatile chemical cues released over a relatively large area that indicate a locale where more specific host cues are most likely to be found. Habitat cues differ from host cues in that they tend to be released in larger quantities, are more easily detectable over longer distances, and may lack specificity, yet provide an effective way for insects to maximise their chances of subsequently encountering specific host cues. This review brings together recent advances in this area, discussing key examples and similarities in strategies used by haematophagous insects, soil-dwelling insects and insects that forage around plants. We also propose and provide evidence for a new theory that general and non-host plant volatiles can be used by foraging herbivores to locate patches of vegetation at a distance in the absence of more specific host cues, explaining some of the many discrepancies between laboratory and field trials that attempt to make use of plant-derived repellents for controlling insect pests.

Waiting with bated breath: opportunistic orientation to human odor in the malaria mosquito, Anopheles gambiae, is modulated by minute changes in carbon dioxide concentration.

Females of the malaria mosquito, Anopheles gambiae, predominantly obtain blood meals within human dwellings. Being highly anthropophilic, human skin odor offers a reliable, host-specific cue, but the challenge posed by pervasive human odor found indoors from used clothing, bedding etc. remains unclear. Anopheles gambiae spends much of its adult life indoors, constantly exposed to human odor even when dwellings are unoccupied. In landing assays, we found that female mosquitoes respond very weakly to human skin odor alone, suggesting that, alone, it is an ineffective landing cue. Landing, however, was dramatically increased by addition of carbon dioxide at a range of concentrations above ambient. Indeed, this effect was seen even when carbon dioxide was just 0.015% above ambient within the assay cage. The synergistic effect of added carbon dioxide quickly waned, thereby facilitating a highly adaptive "sit-and-wait" ambush strategy, wherein females ignore persistent human odor until a living human is present. Unexpectedly, landing rates in the presence of added carbon dioxide were almost as robust during daytime, when An. gambiae has previously been assumed inactive, possibly facilitating opportunistic feeding at times of day when human dwellings are occupied intermittently. We suggest earlier studies that showed strong upwind flight behavior toward human odor alone could, in fact, have been demonstrating orientation toward a human dwelling rather than toward a living human. This new interpretation of how human odors mediate upwind orientation and landing in An. gambiae is discussed.

Avian egg odour encodes information on embryo sex, fertility and development.

Avian chemical communication is a rapidly emerging field, but has been hampered by a critical lack of information on volatile chemicals that communicate ecologically relevant information (semiochemicals). A possible, but as yet unexplored, function of olfaction and chemical communication in birds is in parent-embryo and embryo-embryo communication. Communication between parents and developing embryos may act to mediate parental behaviour, while communication between embryos can control the synchronicity of hatching. Embryonic vocalisations and vibrations have been implicated as a means of communication during the later stages of development but in the early stages, before embryos are capable of independent movement and vocalisation, this is not possible. Here we show that volatiles emitted from developing eggs of Japanese quail (Coturnix japonica) convey information on egg fertility, along with the sex and developmental status of the embryo. Specifically, egg volatiles changed over the course of incubation, differed between fertile and infertile eggs, and were predictive of embryo sex as early as day 1 of incubation. Egg odours therefore have the potential to facilitate parent-embryo and embryo-embryo interactions by allowing the assessment of key measures of embryonic development long before this is possible through other modalities. It also opens up the intriguing possibility that parents may be able to glean further relevant information from egg volatiles, such as the health, viability and heritage of embryos. By determining information conveyed by egg-derived volatiles, we hope to stimulate further investigation into the ecological role of egg odours.

Volatile interaction between undamaged plants affects tritrophic interactions through changed plant volatile emission.

Volatile interactions between unattacked plants can lead to changes in their volatile emissions. Exposure of potato plants to onion plant volatiles results in increased emission of 2 terpenoids, (E)-nerolidol and TMTT. We investigated whether this is detectable by the ladybird Coccinella septempunctata. The odor of onion-exposed potato was significantly more attractive to ladybirds than that of unexposed potato. Further, a synthetic blend mimicking the volatile profile of onion-exposed potato was more attractive than a blend mimicking that of unexposed potato. When presented individually, TMTT was attractive to ladybirds whereas (E)-nerolidol was repellent. Volatile exchange between unattacked plants and consequent increased attractiveness for ladybirds may be a mechanism that contributes to the increased abundance of natural enemies in complex plant habitats.

Volatile exchange between undamaged plants - a new mechanism affecting insect orientation in intercropping.

Changes in plant volatile emission can be induced by exposure to volatiles from neighbouring insect-attacked plants. However, plants are also exposed to volatiles from unattacked neighbours, and the consequences of this have not been explored. We investigated whether volatile exchange between undamaged plants affects volatile emission and plant-insect interaction. Consistently greater quantities of two terpenoids were found in the headspace of potato previously exposed to volatiles from undamaged onion plants identified by mass spectrometry. Using live plants and synthetic blends mimicking exposed and unexposed potato, we tested the olfactory response of winged aphids, Myzus persicae. The altered potato volatile profile deterred aphids in laboratory experiments. Further, we show that growing potato together with onion in the field reduces the abundance of winged, host-seeking aphids. Our study broadens the ecological significance of the phenomenon; volatiles carry not only information on whether or not neighbouring plants are under attack, but also information on the emitter plants themselves. In this way responding plants could obtain information on whether the neighbouring plant is a competitive threat and can accordingly adjust their growth towards it. We interpret this as a response in the process of adaptation towards neighbouring plants. Furthermore, these physiological changes in the responding plants have significant ecological impact, as behaviour of aphids was affected. Since herbivore host plants are potentially under constant exposure to these volatiles, our study has major implications for the understanding of how mechanisms within plant communities affect insects. This knowledge could be used to improve plant protection and increase scientific understanding of communication between plants and its impact on other organisms.

Herbivory by a Phloem-feeding insect inhibits floral volatile production.

There is extensive knowledge on the effects of insect herbivory on volatile emission from vegetative tissue, but little is known about its impact on floral volatiles. We show that herbivory by phloem-feeding aphids inhibits floral volatile emission in white mustard Sinapis alba measured by gas chromatographic analysis of headspace volatiles. The effect of the Brassica specialist aphid Lipaphis erysimi was stronger than the generalist aphid Myzus persicae and feeding by chewing larvae of the moth Plutella xylostella caused no reduction in floral volatile emission. Field observations showed no effect of L. erysimi-mediated floral volatile emission on the total number of flower visits by pollinators. Olfactory bioassays suggested that although two aphid natural enemies could detect aphid inhibition of floral volatiles, their olfactory orientation to infested plants was not disrupted. This is the first demonstration that phloem-feeding herbivory can affect floral volatile emission, and that the outcome of interaction between herbivory and floral chemistry may differ depending on the herbivore's feeding mode and degree of specialisation. The findings provide new insights into interactions between insect herbivores and plant chemistry.

Between plant and diurnal variation in quantities and ratios of volatile compounds emitted by Vicia faba plants.

Ratios of volatile phytochemicals potentially offer a means for insects to recognise their host-plant species. However, for this to occur ratios of volatiles would need to be sufficiently consistent between plants and over time to constitute a host-characteristic cue. In this context we collected headspace samples from Vicia faba plants to determine how consistent ratios of key volatile phytochemicals used in host location by one of its insect pests, the black bean aphid, Aphis fabae, were. These were (E)-2-hexenal, (Z)-3-hexen-1-ol, 1-hexanol, benzaldehyde, 6-methyl-5-hepten-2-one, octanal, (Z)-3-hexen-1-yl acetate, (R)-linalool, methyl salicylate, decanal, undecanal, (E)-caryophyllene, (E)-beta-farnesene, (S)-germacrene D, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene, which had previously been found to be electrophysiologically and behaviourally active to A. fabae. Although the quantities of volatiles produced by V. faba showed large between plant and diurnal variation, correlations between quantities of compounds indicated that the ratios of certain pairs of volatiles were very consistent. This suggests that there is a host-characteristic cue available to A. fabae in the form of ratios of volatiles.

Identification of volatile compounds used in host location by the black bean aphid, Aphis fabae.

Behavioral and electrophysiological responses of winged Aphis fabae to volatiles of faba bean, Vicia faba (var. Sutton dwarf), plants were studied and semiochemicals used in host location were identified. In olfactometer bioassays, aphids spent significantly more time in the region of the olfactometer where V. faba volatiles from an intact plant were present than in control regions with clean air. This response also occurred when an air entrainment sample of a V. faba plant was used as the odor source. Coupled gas chromatography-electroantennography revealed the presence of 16 electrophysiologically active compounds in the air entrainment sample. Fifteen of these were identified as (Z)-3-hexen-1-ol, 1-hexanol, (E)-2-hexenal, benzaldehyde, 6-methyl-5-hepten-2-one, octanal, (Z)-3-hexen-1-yl acetate, (R)-(-)-linalool, methyl salicylate, decanal, undecanal, (E)-caryophyllene, (E)-beta-farnesene, (S)-(-)-germacrene D, and (E,E,)-4,8,12-trimethyl-1,3,7,11-tridecatetraene. An olfactometer response was observed to a 15-component synthetic blend that comprised all identified compounds at the same concentration and ratio as in the natural sample, with the aphids spending significantly more time in the treated regions of the olfactometer where volatiles were present than in the control regions. These data are discussed in the context of insect host location and crop protection.

cis-Jasmone induces Arabidopsis genes that affect the chemical ecology of multitrophic interactions with aphids and their parasitoids.

It is of adaptive value for a plant to prepare its defenses when a threat is detected, and certain plant volatiles associated with insect damage, such as cis-jasmone (CJ), are known to switch-on defense metabolism. We used aphid and aphid parasitoid responses to Arabidopsis thaliana as a model system for studying gene expression and defense chemistry and its impact at different trophic levels. Differential responses to volatiles of induced Arabidopsis occurred for specialist and generalist insects: the generalist aphid, Myzus persicae, was repelled, whereas the specialist, Lipaphis erysimi, was attracted; the generalist aphid parasitoid Aphidius ervi was attracted, but the specialist parasitoid Diaeretiella rapae was not affected. A. ervi also spent longer foraging on induced plants than on untreated ones. Transcriptomic analyses of CJ-induced Arabidopsis plants revealed that a limited number of genes, including a gene for a cytochrome P450, CYP81D11, were strongly up-regulated in the treated plants. We examined transgenic Arabidopsis lines constitutively overexpressing this gene in bioassays and found insect responses similar to those obtained for wild-type plants induced with CJ, indicating the importance of this gene in the CJ-activated defense response. Genes involved in glucosinolate biosynthesis and catabolism are unaffected by CJ and, because these genes relate to interactions with herbivores and parasitoids specific to this family of plants (Brassicaceae), this finding may explain the differences in behavioral response of specialist and generalist insects.

Olfactory recognition of host plants in the absence of host-specific volatile compounds: Host location in the black bean aphid, Aphis fabae.

The black bean aphid, Aphis fabae, responds behaviorally to the odor of its host plant faba bean (Vicia faba) in olfactometer bioassays by spending more time in the treated than control regions. We have shown previously that a blend of fifteen volatile compounds emitted by V. faba elicits the same response as a headspace sample of an intact V. faba plant. Here we report that no single compound within this blend fully accounts for the behavioral response and that the responses to individual compounds are different when in the context of the blend. As none of the compounds are specific to the host, we have hypothesized that A. fabae responds preferentially to the blend of compounds when presented in a species-specific combination of volatiles or in ratios specific to V. faba. Future plans to test which of these two hypotheses pertains to host-seeking Aphis fabae are discussed.