Scorpion Feet Smell: Chemosensory Organs in Arthropods...scorpions by Harald Wolf. Despite being built to sniff with their feet, their chemosensory nervous system has amazing resemblance

Vol 15, No.2 November 2013

E-Nose Pty Ltd

INSIDE

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Scorpion Feet Smell: Chemosensory Organs in Arthropods

Feet that Smell

Allergies to Fragrances

New Beer Flavours

2013 AACSS Abstracts

Upcoming Events

ChemoenseS

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CONTENTS

ChemoSense (ISSN 1442-9098)Web: www.chemosense.net Published by E-Nose Pty LtdP.O. Box 488 Gladesville, NSW Australia 2111Ph. (+61 2) 9209 4083 Fax (+61 2) 9209 4081www.e-nose.info

Production TeamEditor & Advertising:Graham Bell, [email protected] Bell and Associates Pty LtdCentre for ChemoSensory Research

Elke Weiler, [email protected] of Ulm, Germany

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Reproduction of ChemoSense in whole or inpart is not permitted without writtenpermission of the Editor Views expressedherein do not necessarily represent those ofthe Publisher. The Publisher disclaims allresponsibility for any action of any kind takenon the basis of information published herein.

E-Nose Pty Ltd

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2 Editorial

3 Main article on scorpion

olfaction

10 Allergies to Fragrances

14 News: Beer flavour

15 Abstracts of the AACSS

Meeting of 2013

21 Forth-coming events We’re BackGraham BellE-Nose Pty Ltd

If you’re new to reading ChemoSense you’ll find new and interestinginformation from the science of the chemical senses that will delight and informyou. Our writers are leaders in the field and their contributions serve ourinternational community of chemosensory scientists and professionals, as well aspeople in business and industry, who value awareness of new knowledge. Ifyou are an existing reader you’ll know that we have had a 6 month holiday, ourfirst break in publishing in 15 years. Such are the vagaries of cost and effortrequired to produce high quality information. Anyway, we’re back, refreshed,with a new website for back-issues and 50% new editorial personnel: ElkeWeiler, joins me as Co-Editor. Elke is pictured above with her nasalneuroanatomical work which has been used to grace the publications,letterheads and coffee mugs of Northwestern University (USA).

Do your feet smell? Or more specifically, do you smell with your feet? No? Thenyou are probably not an arthropod, and certainly not a chelicerat or scorpion. Inthis issue we bring you a fascinating mini-review on the chemosenses ofscorpions by Harald Wolf. Despite being built to sniff with their feet, theirchemosensory nervous system has amazing resemblance to otherinvertebrates and even to you, our valued mammalian readers.

Can fragrances cause allergies? Despite the molecular mechanismsof causation being poorly understood, allergies triggered by smellsare more common than you might realise. A short article on thesubject by Julia Kahle, will tell you more about how they manifestand what can be done to avoid serious health consequences.

Other news and announcements abound in this new edition ofChemoSense.

[email protected]

Graham Bell and Elke Weiler, Co-Editors of ChemoSense

EDITORIAL

Photo: ATP

Photo: Erich Sommer

Elke WeilerUniversity of Ulm

[email protected]

Cover Image: Hardrurus arizonensisscorpion. The scorpion wasilluminated with UV light during thenight, producing a blue fluorescencethat was used to take the picture(commonly known as "blacklighting"by scorpion afficionados). Total lengthof the animal is about 10 cm.

Virtually all animals possess senses ofsmell and taste, or chemosenses. Thelatter term is more appropriate, since inanimals, such as aquatic species, smell andtaste cannot always be distinguished.Chemoreception is indeed the oldest andperhaps most universal sensory modality,considering that even unicellularprotozoans like paramecia orient withregard to chemical gradients. In higheranimals, chemoreceptors are usuallyarranged at the anterior end of the beastin the form of noses, tentacles orantennae - the latter two includingmechanosensors to probe encounteredobjects by touch. This is a most usefulsituation since it informs the animalimmediately about what is coming its way,or what it has encountered on its path oflocomotion. Equally important is theprobing of food before it is consumed.

Arthropods – insects, crustaceans, spidersand related creatures – are no exceptionhere. The antennae of insects, crayfish andcrabs are located on the head,immediately adjacent to eyes andmouthparts (Fig.1A). The antennae arethe primary chemosensory organs. Manybugs possess highly sensitivechemoreceptors, as exemplified in the

well-studied silk moth Bombyx mori.Physiological measurements havedemonstrated that a single molecule of itssexual pheromone bombycol may besufficient to elicit a sensory signal in themoth brain (Schneider et al., 1968;Kaissling and Priesner, 1970; Kaissling,1986). The silk moth thus rivals in olfactorysensitivity many mammals that are knownto possess a keen sense of smell. Thepolar bear is a celebrity here, renownedfor its ability to localise a food source bysmell from more than 100m distance orunder 1m of snow (Brown, 1993).Sensitivity is not all that counts, though.Distinguishing and memorising smells maybe equally important for an animal’sforaging success. The honeybee is apopular example arthropod here, with itsability to distinguish and learn anappreciable number of often subtlydifferent flower odours, flower shapes andcolours (Menzel and Müller, 1996).

One arthropod group appears to havebeen out of luck in the evolution ofchemosensory organs, however. These arethe chelicerats, that is, scorpions, spidersand their kin. Descending from marineancestors, as has all animal life, thesearthropods conquered terrestrial biotopes


3ChemoSensecont. pg 4

Graham BellE-Nose Pty Ltd

[email protected]

Scorpion Feet Smell: Chemosensory Organs

in ArthropodsHarald Wolf Institute for Neurobiology

University of Ulm

D-89069 Ulm

Germany

[email protected] Elke WeilerUniversity of Ulm

[email protected]

Photo: Berlins Wirtschaftssenator Harald Wolf

4

in the Silurian about 430 million yearsago. At that time, they were alreadyadapted to feeding with an anteriorset of appendages, the set that evolvedinto (chemosensory) antennae in theirforebears: insects and crustaceans.Thus, chelicerats do not possessantennae but instead a pair of clawsor pincers eponymously called chelicers(Fig.1B, C, D).

A note on the evolution of arthropodappendages is useful here. Theappendages on the arthropod body –not just antennae but walking legs,mandibles and associated mouthparts– all derive from the same originallyleg-like limb type that supported both(swimming) locomotion and (filter)feeding in a distant ancestor (Buddand Telford, 2009; Scholtz andEdgecombe, 2006; Hughes andKaufman, 2002). A pair of these limbsoriginally belonged to every segmentof the arthropod body (Table 1). Thattype of appendage decorated anorganism that may have looked

somewhat like a brine shrimp and livedbefore 500 million years ago. And itexplains why, as an arthropod, youcannot have the first set ofappendages as antennae if you earlierdecided to use them as preyprocessing chelicers. This is explainedin some more detail in Table 1.

The original, swimming and filterfeeding arthropod legs have evolvedinto a remarkable variety of limbs, fromretaining filter feeding in barnacles towalking and jumping legs in allwalking arthropods, speed-trapmandibles in some ants that feed onspringtails (Gronenberg, 1995a, b),genital appendages where these arepresent, and poison-injecting chelicers(Fig.1C) – to name just a few.

Chelicers are generally known to formthe unpleasant anterior ends inscorpions and spiders (Fig.1B-D). Theymay be particularly nasty in spiderswhere they form poison-injectingclaws, functionally comparable to thetail stinger of scorpions. Moreunpleasantries concern the fact thatthese appendages do not only injectpoison in spiders to incapacitate theirprey, or a potential predator ifdefensive action is required, but inmost chelicerats, enzymes are injected,too, to digest the prey tissues, allowingthe chelicerate to suck up the resultingnutritious juices, a process aptly calledextra-intestinal digestion. In this way,the chelicers replace the mandibles ofinsects and crustaceans as prey-processing structures, providingenzymatic rather than mechanicalbreak-up of larger food items.Depending on the chelicerat group,the chelicers may also help to pluck

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cont. pg 5

Scorpion Feet Smell: Chemosensory Organs in Arthropodscontinued

Fig 1. Arthropod appendages. Images of aninsect (A, head of a female stag beetle inventral view), a camel spider (B), a tarantulaspider (C), and a scorpion (D) illustrate thedifferent implementations of segmentalappendages as walking legs, feelers, or variousmouth parts. The common origin of all theseappendages is illustrated by the segmentedcharacter of the maxilla in the stag beetle, forexample. The appendages are labelled accordingto the list in Tab.1. Note the (superficial)similarity of the stag beetle antenna (A) to thescorpion pectine in Fig.2A-C, the similarity ofthe (ancestral) camel spider pedipalp to awalking leg (B), the distinctly alteredappearance of the pedipalp in the tarantulaspider (C), and the small chelicer pincer in thescorpion, just in front of the eyes, that helpsplucking apart the prey (D) for extra-intestinaldigestion. Photograph in (C) by courtesy andcopyright of Matthias Wittlinger.


5ChemoSense

away pieces of prey that have beendigested not by poison but byregurgitated gut enzymes. This is true inparticular for scorpions (Fig.1D). Thismode of feeding and extra-intestinaldigestion - perhaps a little repulsive foran educated mammal - has stayed withthe chelicerats ever since, from spidersto scorpions and harvestmen. Onlysome mites and ticks just suck liquidsout of their hosts or prey, since theseare fluid anyway.

With all this feeding frenzy going onwithout a nose or antennae, how dochelicerats examine potential prey? After

all, a chemical sense is useful fororientation and food probing in achelicerat as much as it is in any animal.So without the anterior set ofappendages available to evolve intoantennae, what is the chelicerat solutionto the problem of chemosenses? In fact,there are several. Spiders and camelspiders (Fig.1B) have modified theirsecond pair of appendages (thepedipalps) into antenna-like limbs thatare richly endowed with mechano- andchemoreceptors. They are used to probeobjects both mechanically andchemically, much like the insectantennae are. And they may actuallyresemble antennae morphologically andphysiologically, depending on theparticular chelicerate group.

Scorpions have evolved a dedicated setof appendages for mechano- andchemoreception, too, the scorpion“antennae” (Fig.2). Intriguingly, though,these “antennae” are not located nearthe anterior end of the animal, as inmost animals. They are instead locatedposteriorly, behind the four pairs ofwalking legs (remember, scorpions arerelatives of eight-legged spiders), andthey point downward, towards the floor(Fig.2A-C). Their relationship to legs isstill apparent from three joints (redarrows in Fig.2B, C) that allow somemovement, such as lowering onto thesubstrate. These appendages are calledpectines according to their comb-likeshape. That shape is reminiscent ofinsect antennae that often exhibit comb-like structures (Fig.1A) to maximise thesurface available for the presentation ofodorant receptors: also an obviousreason for the comb-like structure in thescorpion pectines. In male scorpions,these appendages may be equipped

cont. pg 6

Scorpion Feet Smell: Chemosensory Organs in Arthropodscontinued

Table 1. Outline of arthropod appendage evolution. A highly schematised basic arthropod is shown onthe left, with eyes, mouth and identical segmental appendages indicated (see text for a brief descriptionof the ancient filter feeding organism with segmentally repeated similar appendages). The evolutionaryspecialisations of the appendages are listed for chelicerats, insects, and crustaceans for comparison. The1st to the 12th appendages are listed individually, although more than 20 segments may exist. Pleasenote that the mouth, while originally located in the frontal segment without appendages, has movedbackwards in the course of evolution by one or more segments. Many details not shown in this coarseoutline are still under discussion.

continuedScorpion Feet Smell: Chemosensory Organs in Arthropods


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ChemoSense

with more than 140,000 sensory cells,both mechano- and chemoreceptors(Wolf 2008). This number of sensorycells is quite comparable to that inhighly chemosensitive insects like thesilk moth mentioned above. And like ininsects, scorpion males usually possesslarger pectines and more sensillaebecause these are used for findingreceptive females. A female scorpionmay possess only 80,000 sensory cellson her smaller pectines, compared tothose 140,000 in males (Wolf, 2008).

With their unusual position on thescorpion body, the pectines are notused to probe air streams fororientation or to examine prey andfood items, but rather for probing thesubstrate in the context of mating, andperhaps prey or food localisation(Gaffin et al., 1992; Gaffin andBrownell, 1992, 1997). Scorpions thusindeed smell and taste with their feet,or actually with their whole legs,evolved into substrate-orientedantenna-like organs. This should notcome as a surprise considering thatmany arthropods have chemoreceptorson the foot segments of their walkinglegs, including flies and locusts(Roessingh et al., 1997; Newland et al.,2000). Although these contactchemoreceptors have not yet receivedmuch attention, it is evident that suchan arrangement is a useful one. Afterall, flies often walk across their foodand in this way their feet can tell themwhere it tastes best and engagementof the mouthparts is warranted.Similarly in the chelicerats,chemoreceptors on walking legs,chelicerae and pedipalps allow findingand probing of food before it isconsumed (Krapf, 1986).

Even more intriguing than the story ofchemosensory appendages inarthropods is perhaps the similarity ofthese organs’ nerve cell projections inthe central nervous systems of virtuallyall animals (Strausfeld, 2012). Theprojections of the chemosensory nervefibres are all organised into glomeruli,little globular compartments ofnervous tissue that each receive inputfrom one particular subtype ofchemosensory cells (Hildebrand andShepherd, 1997; Maresh et al., 2008;

cont. pg 7

Fig 2. Scorpions, their pectine appendages and pectine neuropils. A scorpion Pandinus imperator isoutlined in ventral view in (A), with the central nervous system indicated in red. (B) shows an enlargedventral view of the pectines, illustrating their comb-like appearance (compare stag beetle antenna inFig.1A); red arrows indicate appendage joints. The photograph in (C) is from a different species,Androctonus australis, with distinctly larger pectines, illustrating inter-species variability. The fused, mainpart of the central nervous system is shown in (D) with the relevant elements labelled; pectine neuropilsare emphasised by dotted shading. (E) and (F) show histological sections (10 micrometers thick) of thepectine neuropil in the frontal plane ((E), cross section) and in the horizontal plane ((F), similar orientationas in (D)); the midline of the nervous system is indicated by arrows. The glomerular structure of the pectineneuropils is visualised by a lipid stain that darkens the dense glomerular neuropil portions (details in (Wolf,2008)). Anterior is to the top of the figure in (A)-(D) and (F); dorsal is to the top in (E).


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Strausfeld, 2012). These glomeruli arethe first stage of chemosensory signalprocessing (Sachse et al., 1999;Firestein, 2001). The respective brainareas in arthropods thus look almostlike miniature versions of the olfactorybulbs in mammals and othervertebrates. The striking similarity ofthe chemosensory projection areas invertebrates, insects, crustaceans andother invertebrates has long engagedneurobiologists. Similarities regardingthe chemosensors associated with theanterior ends of the animals and theirfood intake were sometimes explainedby a possible common origin of suchstructures and their association withfood identification very early in animalevolution. The fact that evenposteriorly located chemosensors inscorpions (and also in some otheranimal groups) conform to theglomerular organisation scheme(Fig.2E, F) supports another morecommonly accepted interpretation(not necessarily to the exclusion of thefirst one, though) (Hansson, 1999;Galizia and Menzel, 2000; Maresh etal. 2008; Zou et al., 2009), as will nowbe described.

It appears that a glomerularorganisation, with each chemosensoryneuron subtype mapping in thecentral nervous system to a smallnumber of glomeruli, or perhaps justan individual glomerulus, is the mostsuitable mode of processingchemosensory information. Selectiveprojection of chemosensory neuronsubtypes into individually definedglomeruli in a one-to-one mannerimplies that the number of glomeruliobserved in a particular animal speciesreflects the number of chemosensory

neuron subtypes in that animal(Vosshall et al., 1999; Galizia andMenzel, 2000; Maresh et al., 2008).

Indeed, this assumption holds for theorganisms studied in sufficient detailto address this question, such as mice,honeybees and fruit flies (Firestein,2001). The house mouse, for example,possesses about 1800 glomeruli in itsleft and right olfactory bulbs,respectively, with each glomerulustype represented twice in each bulb(Royet et al., 1988; Maresh et al.,2008). A corresponding number ofgenes, a few more than 850, codesfor olfactory receptor proteins whichare individually expressed in thedifferent subtypes of olfactoryreceptor neurons in the olfactorymucosa of the nose (Glusman et al.,2000; Maresh et al., 2008). Thehoneybee, a small insect that has tobe strictly conservative with the spaceit can allocate to its brain, has asurprising 160 glomeruli in anolfactory lobe (Galizia and Menzel,2000), and a corresponding numberof olfactory genes (Robertson andWanner, 2006). The above scorpionshave about 100 glomeruli but nothingis known yet regarding their spectrumof chemoreceptor proteins or sensoryneurons.

Humans, much like the house mouseas a well studied fellow-mammal, haveabout 350 functional odour receptorgenes, and about the same numberof silent genes from earlierevolutionary time (Go and Niimura,2008). In general, mammals maypossess well over 1000 genes thatcode for olfactory receptor proteins,which corresponds to some 4% of the

cont. pg 8

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cont. pg 9

mammalian genome being devoted toolfaction (Maresh et al., 2008). This isan appreciable investment into onesensory modality. By comparison, ourvisual system is content with just threereceptor proteins for colour vision(Jacobs, 1996), producing the red,green, and blue visual pigments (plusone protein for scotopic vision at lowlight levels), and comparable numbershold for the other senses. Why is thisexceedingly large number ofchemoreceptor proteins andcorresponding sensory cell typesnecessary in chemoreception? In thevisual system, the whole linearspectrum of visible light can beusefully absorbed by three differentphotopigments with slightlyoverlapping absorption spectra. Colouris extracted by comparing the relativelevels of excitation in the threedifferently tuned photoreceptors,allowing for a smooth coding of huesacross the whole spectrum (Jacobs,1996). The situation is dramaticallydifferent for chemoreception. There isnothing like a linear spectrum ofchemical compounds that an organismencounters. Biologically significantchemicals rather differ in manyaspects, including possession ofalcohol, aldehyde or carboxylic acidgroups, aromatic or aliphatic character,and many more that may all occur incombination. Animals thus had toevolve a set of receptor proteins that

are more or less specific in theirbinding properties for subsets ofchemicals, such as monocyclic aromatsor alcohol groups. And the only usefulway to evaluate such a multi-receptorarray appears to be an initial sorting byreceptor subtype in the glomeruli. Thecentral nervous system may thenassess which pattern of glomeruli, andthus receptor subtypes, are active atall, and among those relative excitationlevels may be compared to characterisea distinctive odour quality.

In summary, then, smelling scorpionfeet have not just informed us aboutthe evolution of chemosensory organsin animals, and in arthropodsspecifically, but they also help todevelop a functional interpretation ofthe ubiquitous glomerular organisationof chemosensory brain areas.

Acknowledgements. Sincere thanks go to Elke Weiler for inspiring and supporting thisreview through critical discussion and many hints regarding interesting literature. SteffenHarzsch was a great cooperation partner in several research projects mentioned here.Matthias Wittlinger allowed me to use his photograph of a Tunisian tarantula spider forFig.2C. Ursula Seifert was a great help in finalising the English text. Part of the researchreported here was funded by SFB 156 and by grants Wo466/6 and Ha 2540/2 of theDeutsche Forschungsgemeinschaft.

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REFERENCES

Brown., G. 1993. The GreatBear Almanac. Lyons &Burford.

Budd, G.E. and Telford, M.J.2009. The origin and evolutionof arthropods. Nature 457:812-817.

Firestein, S. 2001. How theolfactory system makes senseof scents. Nature 413: 211-218.

Gaffin, D.D., Wennstrom, K.L.and Brownell, P.H. 1992.Water detection in the desertsand scorpion, Paruroctonusmesaensis (Scorpionida,Vaejovidae). J. Comp. Physiol.A 170: 623-629.

Gaffin, D.D. and Brownell,P.H. 1997. Response propertiesof chemosensory peg sensillaon the pectines of scorpions.J. Comp. Physiol. A 181: 291-300.

Gaffin, D.D. and Brownell,P.H. 1992. Evidence ofchemical signaling in the sandscorpion, Paruroctonusmesaensis (Scorpionida:Vaejovidae). Ethology 91: 59-69.

Galizia, C.G. and Menzel, R.2000. Odour perception inhoneybees: codinginformation in glomerularpatterns. Curr. Opin.Neurobiol. 10: 504-510.

Glusman, G., Bahar, A,Sharon, D., Pilpel, Y., White, J.and Lancet, D. 2000. Theolfactory receptor genesuperfamily: data mining,classification, andnomenclature. Mamm.Genome 11: 1016-1023.

Go, Y. and Niimura, Y. 2008.Similar numbers but differentrepertoires of olfactoryreceptor genes in humans and

chimpanzees. Mol. Biol. Evol.25: 1897-1907.

Gronenberg, W. 1995a. Thefast mandible strike in thetrap-jaw ant Odontomachus:temporal properties andmorphological characteristics.J. Comp. Physiol. A 176: 391-398.

Gronenberg, W. 1995b. Thefast mandible strike in thetrap-jaw ant Odontomachus:motor control. J. Comp.Physiol. A 176: 399-408.

Hansson, B.S. 1999. InsectOlfaction. Springer Verlag,Berlin, Heidelberg, Germany.

Hildebrand, J.G. andShepherd, G.M. 1997.Mechanisms of olfactorydiscrimination: convergingevidence for commonprinciples across phyla. Annu.Rev. Neurosci. 20: 595-631.

Hughes, C.L. and Kaufman,T.C. 2002. Hox genes and theevolution of the arthropodbody plan. Evol. Dev. 4:459–499.

Jacobs, G.H. 1996. Primatephotopigments and primatecolor vision. Proc. Natl. Acad.Sci. USA 93: 577-581.

Kaissling, K.E. 1986. Chemo-electrical transduction in insectolfactory receptors. Annu. Rev.Neurosci. 9: 121-145.

Kaissling, K.E. and Priesner, E.1970. Die Riechschwelle desSeidenspinners. [Smellthreshold of the silkworm.]Naturwissenschaften 57: 23-28.

Krapf, D. 1986. Contactchemoreception of prey inhunting scorpions (Arachnida:Scorpiones). Zool. Anz. 217:119-129.

Maresh, A., Gil, D.R.,Whitman, M.C. and Greer,C.A. 2008. Principles ofglomerular organization in thehuman olfactory bulb –implications for odorprocessing. PLoS ONE 3:e2640.

Menzel, R. and Müller, U.1996. Learning and memory inhoneybees: from behavior toneural substrates. Annu. Rev.Neurosci. 19: 379-404.

Newland, P.L., Rogers, S.M.,Gaaboub, I. and Matheson, T.2000. Parallel somatotopicmaps of gustatory andmechanosensory neurons inthe central nervous system ofan insect. J. Comp. Neurol.425: 82-96.

Robertson, H.M. and Wanner,K.W. 2006. Thechemoreceptor superfamily inthe honey bee, Apis mellifera:expansion of the odorant, butnot gustatory, receptor family.Genome Res. 16: 1395-1403.

Roessingh, P., Städler, E., Baur,R., Hurter, J. and Ramp, T.1997. Tarsal chemoreceptorsand oviposition behaviour ofthe cabbage root fly (Deliaradicum) sensitive to fractionsand new compounds of host-leaf surface extracts. Physiol.Entomol. 22: 140-148.

Royet, J.P., Souchier, C.,Jourdan, F. and Ploye, H. 1988.Morphometric study of theglomerular population in themouse olfactory bulb:Numerical density and sizedistribution along therostrocaudal axis. J. Comp.Neurol. 270: 559-568.

Sachse, S., Rappert, A. andGalizia, C.G. 1999. The spatialrepresentation of chemical

structures in the antennal lobeof honeybees: steps towardthe olfactory code. Eur. J.Neurosci. 11: 3970-3982.

Schneider, D., Kasang, G. andKaissling, K.E. 1968.Bestimmung der Riechschwellevon Bombyx mori mit Tritium-markiertem Bombykol.[Determination of theolfactory threshold of Bombyxmori with tritium-labelledbombycol.]Naturwissenschaften 55: 395.

Scholtz, G. and Edgecombe,G.D. 2006. The evolution ofarthropod heads: reconcilingmorphological, developmentaland palaeontological evidence.Dev. Genes Evol. 216: 395-415.

Strausfeld, N.J. 2012.Arthropod Brains. Evolution,Functional Elegance andHistorical Significance. TheBelknap Press of HarvardUniversity Press, Cambridge,MA.

Vosshall, L.B., Amrein, H.,Morozov, P.S., Rzhetsky, A.and Axel, R. 1999. A spatialmap of olfactory receptorexpression in the Drosophilaantenna. Cell 96: 725-736.

Wolf, H. 2008. The pectineorgans of the scorpion,Vaejovis spinigerus: structureand (glomerular) centralprojections. Arthropod Struct.Dev. 37: 67-80.

Zou, D.J, Chesler, A. andFirestein, S. 2009. How theolfactory bulb got itsglomeruli: a just so story? Nat.Revs. Neurosci. 10: 611-618.

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Allergy to Fragrances

Julia KahleDeutscher Allergie- und Asthmabund e.V. – DAAB

www.daab.de

41061 Mönchengladbach

Germany


11ChemoSense

Julia KahleEmail: [email protected]

Translated by Elke Weiler, with assistance from Ursula Seifert, University of Ulm

ALLERGY TO FRAGRANCES

Odorants nowadays are pervasive ineveryday life. As ingredients of perfumesand non-food-products such as cosmetics,oils, cleaning agents, detergents and fabricsofteners they appear both as naturalessences as well as synthetic fragrances.

However, odorants are not only pleasant,but can also cause severe health problems.

The first sign of an allergy to a fragranceusually is an allergic contact eczema. Typicalsymptoms range from itching skinerythema, weeping blisters, pruritus andscaly skin to chronic eczema in areas thathad direct contact with the allergenicsubstance. The contact dermatitis is a so-called Type-IV-allergy, where the reaction isT-cell mediated and therefore temporallydelayed. Symptoms appear 48 to 72 hoursafter contact with the allergen.

According to the Information Network ofClinical Centers for Dermatology, about 15-20% of the population in Germany areaffected by contact allergy. Fragrances arethe second most common trigger afternickel, with a prevalence of 11.5%.

DIAGNOSING FRAGRANCE ALLERGY

First there is the patient’s medical history. Iffragrances are suspected to provoke theallergic reaction, an epicutaneous patch test

will identify the trigger substance. Allergyspecialists currently use two definedmixtures of fragrances -“Fragrance Mix I”and “Fragrance Mix II”. The vast majority ofpositive reactions are induced by isoeugenoland oakmoss. Since 2007, 26 individualfragrances can be tested which must bedeclared and specified on product labels ofcosmetics according to the labellingregulation by the European Union. This mayfacilitate the search for the trigger but doesnot necessarily make it easier because thefragrance composition in a product usuallyconsists of several individual substances atdifferent concentrations and each withdifferent and distinct allergenic potential.

TREATMENT OF CONTACT ALLERGY TOFRAGRANCES

Contact allergies cannot be cured. Thetherapy is solely based on the avoidance ofthe trigger substance and treatment of thesymptoms.

To reduce the inflammatory response,externally applied corticosteroid-containingproducts are used. In the acute stageaqueous lotions are preferred; in a chronicstage, lipid containing emulsions are stateof the art to relieve the symptoms of dryskin. In addition, urea and glycerol canincrease the moisture retention of the skin.Beside the anti-inflammatory therapy, it is

continuedAllergy to Frangrances

cont. pg 12

Julia KahleDeutscher Allergie- und Asthmabund e.V. – DAAB

www.daab.de

41061 Mönchengladbach

Germany

12 ChemoSense


particularly important to improve the basic care ofthe affected skin area, to regenerate the defectiveskin barrier.

Indispensible for successful treatment is avoidingthe allergy-provoking substances, because witheach new contact, the symptoms will resurge. Inview of the common use of fragrances in everydayproducts this may be difficult, especially, when theuniversal terms “perfume”, “fragrance”, “aroma” or“flavour” are denoted on the product. Thecomposition and concentrations of the individualingredients are not declared – except for the 26fragrances with a commonly high allergy potential(see above). They must be listed individually on theproduct label with their INCI-name (INCI =international nomenclature of cosmetic ingredients).Declaration is mandatory according to theCosmetics Directives, if the concentration of thesefragrances exceeds 0.01% in products that do notremain on the skin (“rinse-off”, such as shower gelsand shampoos) or is more than 0.001% in productsthat remain on the skin (“leave-on”, for examplelotions, make-up, sunscreen, and deodorants).According to the Detergent Regulation allergenicfragrances have to be marked also in detergents,fabric softeners and cleaning products if theconcentration is above 0.01%.

However, labelling regulations do not yet apply toperfumed candles, incense sticks, scented toys,stationary and air scenting products.

INHALED ODORANT EXPOSURE

Many asthmatics and people with chronicobstructive pulmonary disease (COPD) have healthproblems due to airborne odorants which are used,for example in air-fresheners or are distributed byexcessive perfume use. Experts refer to an “odorintolerance” which may significantly reduce thepatients’ quality of life. Odorants can inducemassive breathing difficulties in these patients andmay even result in an acute asthma attack.

SOME SOURCES OF ALLERGIES TO FRAGRANCES

continuedAllergy to Frangrances

Household products

Fragrant plants and essential oils

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Popular cosmetic fragrances


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Vol 14, No.4 October 2012

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Sensory Insights: Recent researchadvances

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Graham Bell and Associates Pty LtdCentre for ChemoSensory Researchwww.chemosense.info www.e-nose.infoISSN 1442-9098

JJoohhnn PPrreessccoottttTasteMatters Research & Consulting [email protected]

EDITORIAL

CCHHOOOOSSIINNGG TTOO LLIIKKEE OORR LLIIKKIINNGG TTOO CCHHOOOOSSEE??

How do you know what foods you like? Simple. It’s that inner voice, thatwarm feeling, that ….. subjective positive ‘something’ whenever certain foodsare thought about or mentioned or available.Tapping into this subjective state is, at firstsight, relatively straightforward. If I want toknow which food you like or which of severalyou prefer, I can ask you. Sensory andconsumer scientists do this all the time. Youcan be asked to provide a rating of liking fora food, or be asked to rank foods in order ofpreference, or simply choose one food outof several to consume. Each of theseapproaches accesses some aspect of what

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LLeessss tthhaann aa ddeeccaaddee aanndd aa hhaallff aaggoo, no-one but a few Stanford geeks hadheard of Google. Now it is known toalmost every computer user in the world.Google and its many versions (such asGoogle Earth, Google Maps and GoogleScholar) serve many purposes, butGoogle’s primary use remains thedirection of a computer user to crucialsources of information. It is possibly themost important humanitarian innovationsince the internet itself. Google is achannel through which the light ofknowledge is accessed by countlessmillions of people. Happy 14

thbirthday,

Google Inc., born 4th

September 1998.

As that ‘birth” was happening, we weredevising the first issue of ChemoSense.We have since published 56 quarterlyissues and over 500,000 words about theimportant field of chemosensory research.In 2003 we changed from printedhardcopy to electronic-onlycommunication. The cost saving wassignificant, and allowed ChemoSense tosurvive and serve an even greaterreadership.

As the power of the internet grows, wenow face questions of whether it ispossible or even necessary to continue inthe brave new world of free, open-access

BByy GGrraahhaamm BBeellll [email protected]

Reprinted from: http://prescotttastematters.blogspot.com.au/ & www.taste-matters.org

Enter the Blogosphere

Vol 14, No.2 March 2012

E-Nose Pty Ltd

Adventures up the Nose

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AAllaann [email protected]

EDITORIAL

Here is a tale about following your nose. How an innocent scientist interested inthe neurobiology of olfactory function found himself “taking the path lesstravelled” from the friendly fields of olfactory neurogenesis to find himself in themiddle of the scary woods of the neurobiology of brain diseases. The path fromunderstanding the molecular and cellular bases of olfactory sensory neuronregeneration led to the adult stem cells of the human olfactory mucosa, and fromthere to olfactory stem cell as models for schizophrenia, Parkinson’s disease andother less well known diseases affecting the nervous system, Hereditary SpasticParaplegia and Ataxiatelangiectasia. Here iswhat I have learned: science does not travel ina straight line.

NNeeuurrooggeenneessiiss iinn tthhee OOllffaaccttoorryy EEppiitthheelliiuumm

It is received wisdom in the olfactory biologycommunity that the sensory neurons in thenose are replaced throughout life. This was firstobserved in the 1930’s and 1940’s in rabbitsand monkeys, originally through experiments todestroy the olfactory organ as a conduit for thepolio virus to reach the brain(1-3). It was even

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TThhee oobbjjeeccttiivvee ooff CChheemmooSSeennssee is tobring the important discoverieshappening in the scientific field ofthe chemical senses to the world.Our readers include present andfuture leaders of industry andscience. The review published inthis issue meets our objectives andwill delight all readers when theylearn of the amazing journey ofdiscovery taken by Alan MacKay-Sim and his colleagues in recentyears. Their journey “ up thenose” has led into a field ofdiscovery “wider than the sky”.

The first gathering of theAustralasian chemosensorycommunity held in New Zealandtook place in December and wasan acclaimed success. Follow theplans for the next AACSS scientific


National Centre for Adult Stem Cell ResearchEskitis Institute for Cell and Molecular TherapiesGriffith University, Brisbane, Australia

A Nose for Discoveries

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E-Nose Pty Ltd

Emerging evidence supporting fatty acidtaste in humans

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RRuusssseellll [email protected]

EDITORIAL

TThhee sseennssee ooff ttaassttee pprreessuummaabbllyy evolved to inform us about the nutritious or toxicvalue of potential foods. The primary organ responsible for the sense of taste isthe tongue, which contains the biological machinery (taste receptors) to identifynon-volatile chemicals in foods and non-foodswe place in our mouth. Once a food entersthe mouth, the tongue aids manipulation ofthe food, assisting breakdown and distributionthroughout the mouth before swallowing thefood. During this critical period of foodmanipulation the tongue is sampling chemicalsin the food, and when food chemicals activatetaste receptors, signals are sent from the tastereceptors to processing regions of the brain.The signals are decoded by the brain and weperceive the taste of the food, which could be

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A central concern for health is thepart played by fat in what we eat.Fat tends to make food moredelicious. Too much fatconsumption has seriousconsequences for individuals. Inthe less-affluent world too littledietary fat is a concern, butcutting down on fat intake, whilepreserving enjoyment of food is amajor concern in the developedworld. Is dietary fat a hiddenflavour booster, which is perceivedonly by its action on othertastants, or is it a taste in its ownright, a sixth “basic taste”, aftersweet, sour, salty, bitter andumami? Russell Keast addressesthis question and returns answersleaning toward, but notcategorically in favour of fattyacids having a primary or basic


Sensory Science GroupCentre for Physical Activity and Nutrition (CPAN)Deakin UniversityBurwood, Victoria, Australia

Mechanisms of taste perception


E-Nose Pty Ltd

On the Scent of GORDElectronic Nose Profiling and Exhaled BreathCondensate Collection in the Diagnosis ofGastro-Oesophageal Reflux Disease

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EEmmmmaa BBooyyss¹¹,, PPaauull TThhoommaass²² aanndd DDeebboorraahh HH YYaatteess³³

[email protected]

EDITORIAL

GGaassttrroo--ooeessoopphhaaggeeaall rreefflluuxx ddiisseeaassee ((GGOORRDD)) is the most common uppergastrointestinal disorder in the Western world and has been associated withrespiratory conditions including chronic obstructive pulmonary disease (COPD) andasthma. Current diagnostic standards for GORDare lacking in sensitivity and specificity, or areinvasive and time-consuming. Novel methodssuch as the use of exhaled breath condensate(EBC) and electronic nose profiling have thecapacity to improve current diagnostics.Improved diagnosis and treatment of GORDcould improve quality of life for those with thiscommon disorder.

The Montreal Consensus (2006) defines GORDas the presence of troublesome symptoms

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CChheemmiiccaall sseennssiinngg technology ismoving ahead in the field of clinicalmedicine, both human and animal.In this issue, Boys, Thomas and Yatescarry the technology forward in afield where it is badly needed:respiratory disease. Respiratoryailments burden a large percentageof the population and strike downboth young and old. Experimentsreviewed here show that progress isbeing made in rapid diagnosis usingan electronic nose. Withimprovements in hardware anddecision algorithms, the technology isset to achieve reliable early diagnosis,leading to better treatment andtimely, life-saving interventions.

Miss the 2011 AACSS scientificmeeting in New Zealand? Read theabstracts from the meeting, inthis issue �


¹Faculty of Medicine, University of NSW, Sydney, Australia²Respiratory Medicine Department, Prince of Wales Hospital, Sydney, Australia

³Thoracic Medicine Department, St Vincent’s Hospital, Sydney, Australia

Diagnose withArtificial Olfaction

Vol. 11 No.1 December 2008

Editorial

E-Nose Pty Ltd

A Brief ‘Taste’ of the HighEnd Sensory Experience

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Graham Bell and Associates Pty LtdCentre for ChemoSensory Researchwww.chemosensory.comISSN 1442-9098

RRiieekkoo SShhooffuu 11,, MMaarrccoo BBeevvoolloo 22 aanndd HHoowwaarrdd MMoosskkoowwiittzz 33

BBrraannddss aanndd tthheeiirr ‘‘SSeennssoorryy EExxppeerriieennccee’’

Adapting sensory preferences by countries to each product or service is the ordinaryapproach to creating a successful, differentiated product. Yet, when we think about‘brand’, we have to travel deeper. Pursuing ‘personality’ is an eternal theme for a brand.The so-called ‘brand personality’ has to be coherentacross countries. Through experience with the fivesenses, strong brands build a bond to consumerswith a defined, sensorial ‘personality’. If you wantto get a sense of this bond, just think of yourfeelings when you cuddle up someone you love.You feel comfortable with his/her not onlyappearance but also feeling of skin, smell of body,and lovely voice.

Applying our knowledge on the practical, appliedend, we have come to recognize that ‘building’ aclear ‘sensory’ personality is essential requirementof High End, especially for some brands such asWhole Foods, Apple, and Bang & Olufsen. The

Designing Experience

A Taste of Proust

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ChemoSensoryExperience

MMaannyy ggiiffttss ppuurrcchhaasseedd in the comingholidays will be of better thanaverage quality as people seek“premium value” in what they giveand receive. Some people canafford and perhaps cannot resistthe very best, as they perceive it tobe. They are “High End” buyers andwhen they choose brandedproducts they are responding to“High End” brand design. This issueleads with the thoughts of threeexecutives in the business ofcreating global brand images forgoods and services that will fetchthe highest prices in theircategories: so called High Endproducts. They argue that sensoryexperience designed into the“personality” of the brand creates


1 Hakuhodo Inc.Tokyo, [email protected]

2 Philips DesignEindhoven, The [email protected]

3 Moskowitz Jacobs, Inc.White Plains, NY, [email protected]

Vol. 12 No.1 December 2009

Editorial

E-Nose Pty Ltd

Sensory education of schoolchildren: What can beachieved?

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Graham Bell and Associates Pty LtdCentre for ChemoSensory Researchwww.chemosensory.com www.e-nose.infoISSN 1442-9098

SSaarrii MMuussttoonneenn

MMoorree tthhaann ttwweennttyy yyeeaarrss aaggoo, French authors Puisais and Pierre (1987) published the

book Le goût et l’enfant that describes a program to activate school children to using

their senses in food perception. The program

consists of practical exercises, with ten lessons

demonstrating sensory perceptions in different

modalities, their interactions and their

implications. This program was the basis for a

combined effort in which several European

institutions worked to bring a sensory education

program to public schools. There remains

inadequate information of the impact of the

program. In this article we describe our efforts to

scientifically quantify the value and effect of

sensory education.

Learning to taste

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In the 1950s, children in Australiawere given a small bottle of milk inthe classroom, every day, as part ofa post-war national nutritionprogram. The milk had often beenwarmed in the sun outside theclassroom for too long.Nevertheless, the order was thateach child must drink his/her share.The result of enforced ingestion ofsoured milk has left a life-longaversion to drinking milk in some ofthose children. Early sensoryexperience can be the basis for longlasting dietary habits.

On the positive side, as SariMustonen and Hely Tuorila show inthis issue, sensory experiencedelivered in a systematiceducational program, will decreasefear of trying new foods by youngchildren. Sensory education maytherefore provide an important


Department of Food Technology, University of Helsinki, Finland Current affiliation: Valio Ltd, R&D, Helsinki, [email protected]

HHeellyy TTuuoorriillaa

Department of Food Technology, University of Helsinki, [email protected]

www.chemosense.nethttp://www.youtube.com/watch?v=bdLXiCpEVMw

AACSS 2013:Abstracts

NEWSGermans Hopping

for Novel Beer FlavoursBy Graham Bell: [email protected]

14 ChemoSense


BEER WITH MELON FLAVOUR, ANYONE?

A HINT OF PEACH AND PASSIONFRUIT?

Surely the originators of lager and pils would say “nein

danke!”. But no, flavoured beer is taking off in Germany

and it’s all legally brewed according to the “beer purity

law”, thanks to flavour conveyed to the brew through the

hops: naturally flavoured hops.

According to German law, beer can consist only of water,

hops, malt and yeast. No artificial flavour is allowed. But,

when the hops already have a fruity note, you can produce

fruity-beer legally in the home of lager and pils. Bred in the

USA and elsewhere, flavoured hops have been used for

several decades “out there”, even as far away as New

Zealand. Now flavoured hops have been exported to

Germany where beer consumers are taking a shine to the

novel tastes imparted to their favourite beverage

(http://www.dradio.de/dlf/sendungen/umwelt/2253137/).

The term "noble hops" traditionally refers to varieties of

hops which are low in bitterness and high in aroma.

European cultivars include Hallertau, Tettnanger, Spalt, and

Saaz.

The game is on to breed new cultivars with interesting

novel tastes. Where will it lead? Beer tasting of sausage

and sauerkraut, perhaps?

Other references:

http://de.wikipedia.org/wiki/Founders_Organic_Brewery

http://en.wikipedia.org/wiki/Craft_beer#Craft_beer

http://en.wikipedia.org/wiki/Saaz_hops

http://en.wikipedia.org/wiki/Hops

AACSS 2013:Abstracts

1. FUNCTIONAL ANALYSIS OF NEMATODEGPCRS IN YEAST

Muhammad Tehseen, Alisha Anderson,Mira Dumancic, Lyndall Briggs andStephen Trowell.

CSIRO Food Futures Flagship and Division ofEcosystem Sciences, Black MountainLaboratories, Canberra, Australia.

The yeast Saccharomyces cerevisiae is well-known available host for human G-proteincoupled receptor ligand screening due to theease of genetic manipulation, low cost, rapidgrowth, and eukaryotic secretory pathway.Although the Caenorhabditis elegans genomewas sequenced 13 years ago and encodes over1,000 GPCRs, of which several hundred arebelieved to respond to volatile organic ligands,only one of these receptors, ODR-10, has beenlinked to a specific ligand, 2,3-butanedione.Odr-3 is potential G-protein αsubunit involved inodorant detection and activated by Odr-10. Herewe report the functional coupling of the Odr-10to yeast pheromone signalling pathway usingthe yeast - C. elegans chimeric Gα subunit(Gpa1: Odr-3) was used. Interaction betweenOdr-10 with Odr-3 and chimaera was confirmedusing split ubiquitin yeast two-hybrid system.We also report the tailoring of a Saccharomycescerevisiae strain for the analysis of C. elegansolfactory receptor function. In this study, a yeastgpa1� ste2 sst2 far1 quadruple mutantwas used to develop a strain that efficientlycouples a nematode olfactory receptor with theyeast signalling pathway. We used two differentreporter genes: green fluorescent protein (GFP)and LacZ to verify activation of the signaltransduction pathway by ligand-GPCRinteractions. With this heterologouslyengineered yeast system, we will be able toaccelerate the de-orphaning of C. elegans GPCRproteins.

2. TASTE RECPTORS AND FOOD PREFERENCESIN DROSOPHILA

Anupama Arun Dahanukar

U.C. Riverside, USA

Animals rely on their taste systems to selectappropriate foods for consumption. We use themodel insect Drosophila melanogaster tounderstand the mechanisms by which tasteneurons recognize various categories ofchemicals. Sweet and bitter compounds aredetected by members of a large family ofGustatory receptors (Grs), which are expressedin complex combinatorial patterns in tasteneurons. Functional analysis of Grs iscomplicated by the observation that individualtaste neurons express multiple receptors, andalso by the failure to develop a suitable ectopicexpression system. We have developed a novelin vivo expression system to “decode” tastereceptors and have found that individual Grsbelonging to a highly conserved clade of eightGrs are determinants of sweet ligand specificity.Interestingly, we also found that these “sweet”Grs are directly and selectively inhibited bybitter alkaloids, suggesting combinatorialmechanisms for sweet and bitter detection. Wehave recently tested an internal nutrient-sensingreceptor, Gr43a, and its An. gambiae ortholog,AgGr25 in our ectopic expression system andfound that both receptors show robustresponse to fructose and are broadly tuned tosugars. We are now poised to furtherinvestigate mechanisms of Gr function inDrosophila and other insects.

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Anandasankar Ray

PLENARY LECTUREU.C. Riverside, USA

Insects that transmit deadly diseasesto hundreds of millions of people

every year and destroy nearly a thirdof agricultural output utilize theirolfactory system to find hosts. Wehave developed computational andneurophysiological approaches toidentify odorants that modify the

responses of highly conservedolfactory receptors involved in

attraction towards hosts. Using theseodorants we can modify the host-seeking behavior of these deadlyinsects in two ways: "mask" their

attraction to us and "pull" them awayfrom us towards traps. In a secondline of research we have identified

highly conserved receptors thatdetect repellents such as DEET. Usingcomputational and neurophysiology

approaches we have identifiedseveral new repellents with greatly

improved properties. The integratingthe three approaches of "mask", "pull"and "push" provide the foundation for

a new generation of odor-basedstrategies for behavior control agentsthat can have tremendous impact in

the control of disease such as malariaand dengue globally.

Abstracts of the 2013 Scientific Meeting of the Australasian Associationfor ChemoSensory Science (AACSS) held at Phillip Island, Victoria,Australia, 16-18 August 2013.


15ChemoSense

AACSS 2013: Abstracts continued


3. DEVELOPMENT OF A HIGH THROUGHPUTCELL-BASED ASSAY FOR CHARACTERIZINGLEPIDOPTERAN OLFACTORY RECEPTORS

Jacob A. Corcoran1,2, Melissa D. Jordan

1,

Richard D. Newcomb1,2

1. The New Zealand Institute for Plant & FoodResearch Limited, Auckland, New Zealand

2. School of Biological Sciences, University ofAuckland, Auckland, New Zealand

The development of rapid and reliable assays tocharacterize insect odorant (ORs) andpheromone receptors (PRs) remains a challengefor the field. Previously insect ORs and PRs havebeen functionally characterized in vitro throughexpression in Xenopus oocytes, Sf9 cells andHEK293 cells. While these approaches havesucceeded, these systems have inherentcharacteristics that prohibit them from beingused in high throughput formats. We havedeveloped an assay system whereby we canfunctionally characterize insect ORs and PRs in96-well plates using a fluorescentspectrophotometer.

We began by making a T-REx HEK293 cell linecapable of regulating the transcription of genesfrom plasmids containing the ‘tetracyclineoperator’ promoter sequence. After confirmationof Tet-Repressor function, this cell line was stablytransfected with the Epiphyas postvittana OR co-receptor (EposORCO). The cell line was thensingle-cell sorted and resulting clones wereevaluated for inducible EposORCO expression byRT-PCR and western blot and localization byimmunofluorescence. Receptor function was alsoconfirmed by measuring calcium flux in responseto the ORCO agonist, VUAA1. The isogenic cellline with the best inducible expression andfunction of EposORCO was chosen forsubsequent expression and characterization of E.postvittana ORs and PRs.

To date we have identified 72 putative olfactoryreceptors from genome and transcriptomelibraries of E. postvittana. Quantitative RT-PCRanalyses have revealed a subset of these genesthat display male-biased antennal expression,making them likely candidates for being E.postvittana PRs. Here we report theresponsiveness of one of the male-biased E.postvittana receptors to the moth’s sexpheromone components to demonstrate theutility of the system.

4. LIGAND-BINDING OF THREE PHEROMONE-BINDING PROTEINS IN THE BEETARMYWORM SPODOPTERA EXIGUA

Nai-Yong Liu1,2, Fang Yang

1, Alisha

Anderson2, Wei Xu

2, Shuang-Lin Dong

1*

1Education Ministry Key Laboratory of IntegratedManagement of Crop Diseases and Pests,College of Plant Protection, Nanjing AgriculturalUniversity, Nanjing 210095, China,

2CSIRO

Ecosystem Sciences, Black Mountain, AustralianCapital Territory 2601, Australia

Moth pheromone-binding proteins (PBPs) playcrucial roles in pheromone reception. InSpodoptera exigua, three SexigPBPs (PBP1-3)were cloned from the antenna, and are classifiedinto three distinct sub-groups. Considering thesex pheromones of S. exigua have beenidentified as a four-component blend, it ishypothesized that each PBP may be tuned to aspecific component (s) of the blend. To test thishypothesis, we carried out ligand-binding assaysof three SexigPBPs to sex pheromones,pheromone analogs and plant volatiles. Ourresults show all three SexigPBPs can bind alltested sex pheromones, suggesting theseSexigPBPs have no obvious discrimination amongdifferent pheromone components. However,SexigPBP1 exhibited much stronger bindingaffinities to sex pheromones and their analogsthan the other two SexigPBPs (PBP1 >> PBP2 >PBP3), especially those ligands with a doublebond in the 9th position. Plant volatiles provedto be poor ligands for all three SexigPBPs. Inaddition, binding of SexigPBP1 and SexigPBP2 tothe main sex pheromone Z9,E12-14:Ac werestrongly affected by pH, in contrast bindingaffinity of SexigPBP3 appeared to be only slightlyaffected. Similar results were also observed in itssibling species S. litura. Our results suggest thatSexigPBP1 might play major roles in the processof female sex pheromone reception.* Corresponding author: Shuang-Lin DONG,[email protected].

This work was supported by grants fromNational Natural Science Foundation (31071978)and the Agricultural Ministry (201203036) ofChina.

5. SENSING SIMILARITIES: DO PIGS TASTE LIKEHUMANS?

Nadia de Jager1and Eugeni Roura

1

1The University of Queensland, Centre forNutrition and Food Sciences (QAAFI)

The exploration of the complexity of the tastesystem is gaining momentum, particularly inlight of its role in nutrient sensing. The diverselocalisation of the expression of taste receptor(TR) genes in oral and in non-oral tissues, suchas in the gastrointestinal tract, has beenestablished, predominantly as a result of rodentresearch. However, rodents and humans areoften quite dissimilar in terms of their dietaryhabits and digestive physiology. In contrast, thepig has gained status as a main biomedical andnutritional model for humans. Our primaryobjectives are firstly to catalogue and study theTR repertoire; and secondly to explore the notionthat this system is involved in mediating thenutritional status and appetite in pigs. Ourresults from quantitative real time PCR assaysshow that the pig and the human TR repertoireshare high similitude, except when consideringthe bitter family (Tas2R). For example, whilehumans have 25 Tas2Rs, the pig only has 16.Expression of the genes encoding sweet andumami (Tas1R1, Tas1R2, Tas1R3), fatty acid(GPR120, GPR40, GPR41, GPR43 and GPR84),glutamic/amino acid (mGLUR1, mGLUR4,GPRC6A, and GPR92) sensing receptors have allbeen identified in pig tissues as a result of ourresearch. With the exception of Tas1R2, which isnot always abundantly expressed, the tissues inwhich we have confirmed TR expression includetongue, stomach ridge and -antrum, duodenum,jejunum, ileum, colon proximal and distal,caecum and liver. In addition, we have studiedthe impact of nutritional interventions onchanges in the level of TR expression andpreliminary results will be discussed. Overall, ourresults support the view that pigs have a well-developed taste system and, with the exceptionof the bitter taste family, that a high similarity isshared with humans.

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16 ChemoSense



6. SPECIES-SPECIFIC SETS OF OLFACTORYRECEPTOR NEURONS IN INSECTS

Kye Chung Park (New Zealand Institute forPlant and Food Research)

Insects exhibit highly sensitive and selectiveperipheral olfactory sensory system. Theolfactory sensillum, an independent unit housingone or a few olfactory receptor neurons (ORNs),is the key element of detecting external chemicalsignals, and delivers the information to the brainfor further processing. Here we show theelectrophysiological response profiles of ORNs insix different insect species (four species of mothsand two species of weevils) to a panel of theirhost and non-host plant volatile compounds.Many of the ORNs were highly specialized fordetecting a narrow range of volatile compoundsalthough some ORNs showed broad spectrumresponses. The profiles of ORNs were species-specific, and the differences in the ORN profilesappeared to be greater between the twodifferent taxonomic groups (moths vs. weevils)than those between species within the sametaxonomic group. The ORNs and theircorresponding active volatile compounds in eachspecies could be correlated to its host and non-host plants. Each species appeared to have twodistinct sets of ORNs, one specialized for thehost-plant specific volatile compounds and theother for non-host plant specific volatilecompounds. It is suggested that thesephatophagous insects use the combinationalinput from these two sets of ORNs to locatetheir host plants and discriminate them fromnon-host plants.

Key words: electrophysiology, host plant, insect,olfaction, olfactory receptor neuron, sensilla,volatile compounds

7. PHENOTYYPIC EXPRESSION OF OLFACORYRELATED GENES IN SEXULAS ANDWORKERS OF THE LEAF-CUTTING ANTA.VOLLENWEIDERI

Sarah I Koch1,2Christoph J Kleineidam

1, Bill

S Hansson2, Ewald Grosse-Wilde

2

1University of Konstanz, Department of Biology,Universitätsstraße 10, 78464 Konstanz, Germany2Max Planck Institute for Chemical Ecology,Beutenberg Campus, Hans-Knöll-Straße 8, 07745Jena, Germany

Leaf-cutting ants are highly derived eusocialinsects, having enormously large colony sizes.Their division of labor is mainly based onolfactory communication, where different sizedworkers perform distinct odor-guided behaviors[1]. Workers and sexuals (queens and males) ofA. vollenweideri were found to exhibit differentantennal lobe (AL) phenotypes, indicatingdifferences in odor information processing [2,3].For example large workers have one extremelylarge glomerulus in the AL (macroglomerulus,MG), and it is associated with trail-pheromonedetection [2]. ALs fo males contain three MGswhich most likely are involved in sex-pheromonedetection. We investigate the molecular basis ofthe developmentally induced plasticity in A.vollenweideri by analyzing antennal transcriptsand phenotype-specific microarrays. Within theantennal dataset we identified 70 odorant-receptor coding genes (OR), as well as theortholog of the Olfactory Co-receptor (Orco).Analyzing the microarray data revealed that oneOR is highly expressed in large workerscompared to tiny workers (relative expression>3). In males, two ORs are highly expressed inthe antenna compared to queens. Due to theone glomerulus - one receptor wiring in the AL[4], high expression levels of ORs are likely tocorrelate with glomerular size, making thempotential trail and

sex pheromone receptors. Most of the ionotropicreceptor coding genes identified in A.vollenweideri are highly expressed in malescompared to queens. Odorant binding proteincoding genes (OBPs) and chemosensory proteincoding genes (CSPs) show no consistentdifferential expression pattern across the workersubcastes or the sexuals.

Funded by DFG: SPP 1392

8. A PHOSPHOLIPID FLIPPASE REQUIRED FOROLFACTORY RECEPTOR NEURON FUNCTIONIN DROSOPHILA MELANOGASTER

Michelle Pearce, Yu-Chi Liu, TakahiroHonda, Marien deBruyne, Coral G. Warr

School of Biological Sciences, Monash University,VIC, Australia

Drosophila olfactory perception is initiated whena chemical ligand from the environment binds toan odorant receptor (OR) localised to thedendrites of odorant receptor neurons (ORNs). Ina search for new olfactory genes, we undertooka large scale EMS-induced mutant screen usingelectrophysiology to test for olfactory defects.We found a single mutant with a pronouncedreduction in olfactory response to all odorantstested, with the exception of CO2. Deficiencymapping identified a region consisting of twelvecandidate genes of which one, dATP8B, wasidentified as containing a nonsense mutation inthe mutant strain using whole genome re-sequencing. A complementation test confirmedmutations in this gene as the cause of theolfactory defect. RT-PCR and RNAi analysesindicate that dATP8B is expressed in a tissuespecific manner, and is required in ORNsexpressing ORs respectively. This gene encodes aphospholipid flippase, a protein subfamilythought to be essential for maintainingasymmetry of phospholipids in some lipidmembranes, whereby phosphatidylserine andphosphatidylethanolamine are enriched in thecytosolic leaflet. As an otherwise uncharacterisedgene, dATP8B has not been implicated in othercellular processes, and no other phospholipidflippase has been implicated in olfaction. Futurework will include an investigation of the sub-cellular location of this flippase, and analyses ofthe other five Drosophila genes belonging to thisfamily to determine their roles in olfaction and innervous system function.

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18 ChemoSense




9. MOLECULAR BASES OF OLFACTION ANDOLFACTORY PLASTICITY IN LEPIDOPTERA:TRANSCRIPTOMIC APPROACHES IN THECOTTON LEAFWORM SPODOPTERALITTORALIS

Emmanuelle Jacquin-Joly1, Aurore Gallot

1-2,

Arthur de Fouchier1-3, Erwan Poivet

1,

Fabrice Legeai2, Nicolas Montagné

3

1INRA, UMR 1272 INRA-UPMC Physiologie del’Insecte : Signalisation et Communication, routede Saint-Cyr, Versailles, France 21IRISA, équipe Symbiose, Campus universitairede Beaulieu, Rennes, France3UPMC – Université Paris 6, UMR-A 1272Physiologie de l’Insecte : Signalisation etCommunication, Paris, France

The sense of smell is a critical determinant ofvital insect behaviours, including mate and foodseeking, oviposition and predator avoidance. Wehave established the cotton leafworm,Spodoptera littoralis, as a nocturnal insect modelto decipher the underlying mechanisms. Wedeveloped a transcriptomic approach: 1) todescribe a large repertoire of chemosensorygenes in both adults and larvae, 2) to addressolfactory plasticity via digital gene-expressionprofiling.

Sanger and next generation sequencingstrategies allowed us annotating ~ 77000expressed contigs. Among them, we described alarge repertoire of candidate odorant-bindingand chemosensory proteins, ionotropic receptorsand olfactory receptors, these latter beingcurrently deorphanized via expression in theDrosophila empty neuron system coupled withsingle neuron electrophysiological recordings.Comparison between adults and larvae revealeddifferent but somewhat overlapping expressionof the chemosensory genes in the differentdevelopmental stages.

The transcriptome was also used to investigatethe transcriptional changes induced by starvationin the chemosensory tissues of S. littoraliscaterpillars, using RNAseq expression profiling.~4 million short reads obtained from antennaeand maxillary palps dissected from fed and 24 hstarved larvae were mapped on the referencetranscriptome and counted. We revealed bothup- and down-regulated transcripts uponstarvation, including genes potentially involved inolfaction.

These approaches not only establish the use oftranscriptomic sequencing for the identificationof divergent chemosensory receptors in a speciesfor which no genomic data are available, butalso enable investigation of chemosensorymodulation via digital gene-expression profiling.

10. ANATOMICAL DEVELOPMENT OF THEHUMAN SENSE OF TASTE: COMPLETE BYMID-CHILDHOOD.

David G Laing1and Maryam Correa

2

1 School of Women’s and Children’s Health,Paediatric Discipline, Faculty of Medicine, UNSW,Level 3, Sydney Children’s Hospital, High Street,Randwick, NSW, 20312School of Science and Health, University ofWestern Sydney, Locked Bag 1797, PenrithSouth, NSW 1797

During the past decade Laing and colleagueshave shown that the anterior region of thehuman tongue which is rich in taste-sensitivefungiform papillae (FP) and receptor cells, ceasesto grow in size by 8-10 years of age. In contrast,the posterior region, largely devoid of FP, ceasesgrowth at 15-16 years. Importantly, the similarsize of the anterior tongue of children and adultsallows a direct comparison of FP density whichreflects taste sensitivity. Accordingly, the presentstudy with 30 adults and 85 7-12 year olds,aimed to determine the age at which FP density,hence sensitivity, stabilizes, and whether FPdensity in a small region of the anterior tonguecan be used to predict total FP density/numbers.A digital camera was used to photograph thetongue and FP, whilst Adobe Photoshopsoftware allowed analysis of the data. Theresults indicated that the number of papillaestabilized at 9-10 years of age, whilst thedistribution and growth of FP stabilized at 11-12years. Importantly, a single sub-population of FPpredicted the density of FP on the wholeanterior tongue of 7-10 year olds, whereasanother was the best predictor for the olderchildren and adults. Overall, the population, sizeand distribution of FP stabilized by 11-12 years ofage which is very close to the age that cessationof the growth of the anterior tongue occurs.Clinically, the procedure has been used as a non-invasive assessment of taste loss during cancer,chronic kidney disease, and otitis media.

11. THE NOSE TELLS US WHAT TO SEE:MODULATING EFFECT OF ODOURS ONVISUAL PERCEPTION

Amanda Robinson, Jason B. Mattingleyand Judith Reinhard

The University of Queensland, Queensland BrainInstitute, St Lucia QLD 4072, Australia

Multisensory integration is an inherent feature ofperception yet the interactions between olfactionand vision are not well understood. While it isknown that vision can influence how we perceivecertain odours, few studies have providedconclusive evidence that odours can impact onthe visual sense. This study investigated whetherodours influence selective attention towardsfamiliar visual objects during the attentional blink(AB), a phenomenon where the second of twotargets in a rapid serial visual presentationstream (RSVP) is undetected if it appears 200-500ms after the first target, due to a bottleneckin the temporal allocation of selective attention.Participants monitored a rapid stream of colourphotographs of objects to discriminate whichvisual object with a characteristic smell (T2;lemon, orange, rose or mint) appeared after aninitial target (T1). During the task, participantswere exposed to an odour that was congruent,incongruent or irrelevant with respect to T2. Wefound that a congruent odour significantlyenhanced the salience of the odour-relatedimage and attenuated the attentional blinkcompared to an incongruent or irrelevant odour.This effect was not observed when the odourcue was replaced by a word cue, demonstratingthat it was a sensory and not semantic effect.Indeed, a follow-up EEG study showed thatcongruent odours significantly enhance earlyneural responses to matching visual objects,implying an important influence of olfactoryinputs on early visual responses. Our resultsprovide evidence that our visual perception issignificantly influenced by our sense of smell.

cont. pg 19


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12. OSTRINIA NUBILALIS STRAIN SPECIFICPHEROMONE PERCEPTION: FIRSTPHYSIOLOGICAL AND MOLECULAR MAP

F. A. Koutroumpa1,2,3

, Z. Kárpáti1,4, C.

Monsempes3, S. Hill

1, B.S. Hansson

5, E.

Jacquin-Joly3, J. Krieger

6, T. Dekker

1

1Swedish University of Agricultural Sciences,Department of Plant Protection Biology, Divisionof Chemical Ecology, PO Box 102,Växtskyddsvägen 3, SE-230 53 Alnarp, Sweden.2Max Planck Institute for Chemical Ecology,Department of Entomology, BeutenbergCampus, Hans-Knöll str. 8, 07745 Jena, Germany.3INRA, UMR PISC Physiologie de l'insecte, INRA,Route de Saint-Cyr, 78026 Versailles cedex,France.4Hungarian Academy of Sciences, Centre forAgricultural Research, Plant Protection Institute,Department of Zoology, Budapest, Pf 102 H-1525, Hungary.5Max Planck Institute for Chemical Ecology,Department of Neuroethology, BeutenbergCampus, Hans-Knöll str. 8, 07745 Jena, Germany.6University of Hohenheim, Institute ofPhysiology, Garben str. 30, 70599 Stuttgart,Germany.

Moth pheromone communication is particularlyillustrative in the study of evolution of signals forspecies recognition and subsequent prematingisolation. We searched for the mechanism thatcaused Ostrinia nubilalis to display a dimorphismin pheromone preference. More precisely howdoes the behavioral preference and physiologicaldifferences between strains correlate with themolecular 'code' of the olfactory system? Wecombined whole mount double in situhybridization and quantitative PCR on the sevenpheromone-receptors of O. nubilalis, with singlesensillum recordings on sensilla trichoidea. Wefound only one sensillum type housing sensoryneurons tuned to pheromone. This sensillumcontained three sensory neurons detecting thetwo pheromone components (Z11-14:OAc andE11-14:OAc) and the behavioral antagonist (Z9-14:OAc), respectively. We demonstrate thatparental strains and hybrids differ consistently inthe specificity of the 3 co-housed neurons, withtranscripts’ expression differences andreshuffling of sensory neuron size, not onlybetween the E11 and Z11 neuron but among allneurons that co-inhabit this sensillum. We alsodemonstrate that sensory neurons mediatingbehavioral antagonism may co-express anunusually high number of receptors, conferringbroad tuning to behavioral antagonist. This

elegantly allows the ‘olfactory code’ to keep itssimplest tripartite setup, while not affectingolfactory precision. Finally, we found an unusualdichotomous receptor expression in a singlesensillum type, indicative of an intermediate stepin a sensillar split. These findings are ofimportance in understanding coding andevolution of pheromone preference andantagonism.

13. CHEMOSENSORY GENES FROM COTTONBOLLWORM HELICOVERPA ARMIGERA

Wei Xu, Alexie Papanicolaou and AlishaAnderson

CSIRO Ecosystem Sciences, Black Mountain,ACT, Australia 2601

The chemosensory system is critical in guidinginsect behaviours. Helicoverpa armigera(Hübner) is one of the most polyphagous pestspecies in the world, so studies on itschemosensory system may give insights into themolecular mechanisms that underlie its ability toexploit a wide variety of host crops and helpdevelop new approaches to control it. Wesequenced and assembled transcriptomes fromlarvae antennae, larvae mouthparts, adult heads,adult tarsi and female adult abdomen. Weidentified fifty-eight odorant receptors (ORs),nine gustatory receptors (GRs), thirty-eightodorant binding proteins (OBPs) and nineteenchemosensory proteins (CSPs). The expressionpattern of these genes were also analysedamong different tissues, ages and genders.Seven candidate pheromone receptor (PR) geneswere analysed by calcium imaging analysis andHarmOR13 showed significant responses to themajor sex pheromone component, (z)-11-hexadecenal. This study will improve ourunderstanding of the insect chemosensorysystem and assist in the development of moreenvironmentally friendly, pheromone basedcontrol strategies.

cont. pg 20

20 ChemoSense



14. SEX PHEROMONE EVOLUTION ANDSPECIATION IN THE NEW ZEALANDENDEMIC LEAFROLLER MOTHS OF THEGENERA CTENOPSEUSTIS ANDPLANOTORTRIX: THE MALE’S STORY

Bernd STEINWENDER1,2,3, Thomas

BUCKLEY3,4, Richard D. NEWCOMB

1,2,3

1 The New Zealand Institute for Plant & FoodResearch Limited, Auckland, New Zealand2 School of Biological Sciences, University ofAuckland, Auckland, New Zealand3 Allan Wilson Centre for Molecular Ecology andEvolution4 Landcare Research, Auckland, New Zealand

The major feature of mate recognition in mothsis the ability of the male to distinguish betweensex pheromone blends of their own and otherspecies. While we are beginning to understandthe molecular basis for differences betweenspecies of moths in their ability to producedistinct pheromone blends, little is known ofhow the males’ pheromone reception systemevolves to track any newly evolved pheromoneblend, especially at the molecular level. Toaddress this question we are studying siblingspecies pairs within the New Zealand endemicleafroller genera Ctenopseustis and Planotortrix.To identify putative pheromone receptorspreliminary genome assembles and antennaltranscriptomes of C. obliquana and P. octo weresearched and odorant receptor genes tested formale-biased expression in their antennae byquantitative RT-PCR. To date, five candidatepheromone receptors from C. obliquana and C.herana and three from P. octo and P. excessanahave been identified. Functional assays revealthat OR07 from the Ctenopseustis speciesresponds to (Z)-8-tetradecenyl acetate. Currentwork involves further functionally testingcandidates in cell-based assays for the ability todetect sex pheromone components and makingcomparisons between species both in receptorsequence and gene expression levels to identifythe differences between species in the male’sside of mate recognition.

15. A MENDELIAN TRAIT FOR OLFACTORYSENSITIVITY AFFECTS ODOR EXPERIENCEAND FOOD SELECTION

Sara R. Jaeger1, Jeremy F. McRae

1,

Christina M. Bava1, Michelle K. Beresford

1,

Denise Hunter1, Yilin Jia

1, Sok Leang

Chheang1, David Jin

1, Mei Peng

1, Joanna C.

Gamble1, Kelly R. Atkinson

1, Lauren G.

Axten1, Amy G. Paisley

1, Leah Tooman

1,

Benedicte Pineau1, Simon A. Rouse

1,

Richard D. Newcomb1,2,3

.1.The New Zealand Institute for Plant & FoodResearch Limited, Auckland, New Zealand2.School of Biological Sciences, University ofAuckland, Auckland, New Zealand3 .Allan Wilson Centre for Molecular Ecology andEvolution, New Zealand

Humans vary in sensory acuity to many odours.However, the importance of genetic variationand how such variation also affects odorexperience and food selection remains uncertain,given that such effects have been shown inbitter taste perception. Using a genome wideassociation approach we identified major loci forsensitivity to four odours (2-heptanone,isobutyraldehyde, βdamascenone and βionone),each located at or near clusters of olfactoryreceptor (OR) genes. Subsequently we focusedon βionone, which shows extreme sensitivitydifferences. βionone is a key aroma in foods andbeverages, and is added to products in order togive a pleasant floral note. Genome-wide and invitro assays demonstrate rs6591536 as the causalvariant for βionone odor sensitivity. Rs6591536encodes a N183D substitution in the secondextracellular loop of OR5A1, and explains >96%of the observed phenotypic variation, resemblinga monogenic Mendelian trait. Individualscarrying genotypes for βionone sensitivity canmore easily differentiate between food andbeverage stimuli with and without addedβionone. Sensitive individuals typically describeβionone in foods and beverages as ‘fragrant’and ‘floral’, whereas less sensitive individualsdescribe these stimuli differently. Rs6591536genotype also influences emotional associations,and explains differences in food and productchoices. These studies demonstrate that an ORvariant that influences olfactory sensitivity canaffect how people experience and respond tofoods, beverages and other products.

16. NEUROETHOLOGICAL BACKGROUND OFTHE EUROPEAN CORN BORER PHEROMONESYSTEM

Zsolt Karpati

The European corn borer moth (ECB, Ostrinianubilalis; Pyralidae) is a well-known pest of thecorn in the northern hemisphere. Beside its peststatus the species has a pheromonepolymorphism system. Two races exist which useopposite ratio of the two pheromonecomponents causing reproductive isolation innature.

We studied how the difference in malepheromone preference correlates withdifferences in wiring of olfactory input andoutput neurons in the primary olfactory neuropil,the antennal lobe (AL). Activity dependentneuronal staining, intracellular recording andimmunocytochemistry were used to establish thestructure and function of male olfactory receptorneurons (ORNs) and AL projection neurons (PNs).Both races have an indistinguishable pheromonesensitive macroglomerular complex (MGC) but areversed topology. Apparently, the single-gene-mediated shift that causes a radical change inbehavior is located upstream of the antennallobes, i.e. at the ORN level. However, hybrids ofthe two races prefer intermediate ratios. How atopological arrangement of two glomeruli canaccommodate for an intermediate preferencewas unclear. PN recordings and stainings inhybrids show a dominance of the E-type MGCtopology, but volumetric measurements of theMGC and antennal response shows anintermediate preference.

We also studied how the response specificity ofthe males changing during the orientation flightusing a wind tunnel behavioral assay. Males thathave ‘locked on’ to a natural ratio pheromoneplume readily continue plume following alongoff-ratio blends. This implies that malesgeneralize pheromone quality after initialcontact. The relaxation of specificity observedhere may also help explain the substantial andunexpected rates of field hybridization betweenraces. At high population densities, males thathave first encountered filaments of pheromonefrom their own race may stray into a plume fromthe other race and then hybridize.

17 – 20 November 2013 Urban Environmental PollutionBeijing, Chinawww.uepconference.com/index.html

19 – 21 November 2013 Air Quality Measurement - Methods and TechnologySacramento, Californiawww.measurements.awma.org

28 – 31 January 2014 Australian Neuroscience Society (ANS)Adelaide, South Australiawww.ans.org.au

9 – 14 April 2014 AChemSAssociation for Chemoreception SciencesBonita Springs, Florida, USAwww.achems.org

31 May – 3 June 2014 Odors and Air Pollutants ConferenceMiami Floridawww.wef.org/odorsair

30 July – 1 August 2014 The Sensometric SocietyChicago, USAwww.sensometric.org

7-11 September 2014 ECRO: European Chemoreception OrganisationDijon, Francewww.ecro-online.com

18 – 21 March 2015 German Neuroscience SocietyGoettingen, Germanyhttp://nwg.glia.mdc-berlin.de/en/conference/

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