Springer Handbook of Odor

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HHandbookSpringer

of OdorAndrea Buettner (Ed.)

With 458 Figures and 122 Tables

K

EditorAndrea BuettnerFraunhofer Institute for Process Engineering and PackagingGiggenhauser Str. 3585354, Freising, Germanyandrea.buettner@ivv.fraunhofer.de

ISBN: 978-3-319-26930-6 e-ISBN: 978-3-319-26932-0DOI 10.1007/978-3-319-26932-0Springer Dordrecht Heidelberg London New York

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V

Foreword

More knowledge is needed!(Wir können wissen, wir müssen wissen)

Around odors there is a wealth of very differentjobs, but all are wonderful when they are done with pas-sion. Passion is the common denominator for all threeof us, even if we have different activities. What activi-ties, by the way? Some analysis is needed first in orderto see more clearly, so that we can identify more pre-cisely what has to be done for the odor realm.

The three of us agree: Rembrandt was an artist andhe was not a house painter; on the other hand, whendoing his job, a house painter does not look for emotion,and he or she has to be a technician, even if there issome artistic component involved. The most ineffableRembrandt painting is appealing.

We don’t say that technique is better than art, orthat art is better than technique, because the evalua-tion criteria are not the same in the two fields. Shouldwe personally prefer art to technique, as is frequentlyproposed? Probably not, because sometimes we needto paint walls, and Rembrandt is useless in this regard,and sometimes we need something different, and Rem-brandt becomes interesting.

All this shows that our choices have to be carefullyanalyzed, and this discussion is particularly importantin the realm of odors, because it is a place where prefer-ences are ubiquitous. Of course, it is easy to understandthat It smells good indeed means Personally, I like thisodor, so that we can ask the question of the legitimacyof a personal preference, even if this comes from a mas-ter perfumer. How is his preference more important thanthat of a non-specialist? This is a first question . . . ,which we propose not to answer. Our goal here will bethis one: ask questions, because we would love you tothink about them and propose your own answers.

Coming back to jobs, the vision proposed here con-sists in considering that there are different jobs, in theareas of art, technique, technology, science. In order toavoid hierarchy, let’s consider them in alphabetical or-der: art, science, technique, and technology.

Some people, such as Maurice Roucel, with odors,or the French chef Pierre Gagnaire, with cooking, are re-luctant to call themselves artists, because they fear pre-tentiousness, but facts are always themost important. Aswe said before, Rembrandt was not a house painter, andhis project was emotion, even if it included, of course,mastering the technique, and a commercial component.

In art, there is probably a passion for beauty, asPierre Kurzenne says, and much apprenticeship work

Pierre KurzenneBeauty Care &New Molecules Develop-ment Senior PerfumerSymrise, Clichy, France

Maurice RoucelMaster PerfumerSymrise, Paris, France

Hervé ThisDirector AgroParisTech-Inra International Centreof Molecular GastronomyParis, France

is needed, because work never endsin the quest for perfection in art.We slog and we work. Passion fre-quently means being able to see theintimate interest of the work. Butwith respect to beauty, the issue ishuge.

In the kitchen, the issue is not tolook at dishes, but to eat them. Heregood means beautiful to eat; withmusic, it means beautiful to hear(we don’t care if the pianist is welldressed or not). And one can easilyunderstand that with odors, the issueis to make beautiful odors as well,which means odors that we admire.

Beauty is, then, the main ques-tion in art, and it is interesting toobserve that it has had thousands ofdefinitions in the past centuries, butit always had something do to withemotion and culture. This holds forodors; the project of the odor artistcan be to please, to make happy, toanger, to seduce . . . , but never toleave you indifferent.

For Plato, art was bad, becauserepresenting (reproducing the odorof mango, of olive oil, . . . , etc.)meant a double detachment fromthe truth, but at the same time, thephilosopher could not escape dis-cussing art and he had to tackledifficult questions, such as the fas-cination for perfect representation,like the painting of grapes that ap-peared so true that birds were con-fused!

Anyway, Plato was refuted byAristotle! The odor of the raspberrydoes not exist: any particular rasp-berry has a particular odor, and thismeans that the production of theraspberry odor is a lost battle. Wecan only produce one chosen odoramong an infinity thereof. This isthe old debate of realism againstidealism.

Which particular odor of rasp-berry does the perfumer or the fla-

VI Foreword

vorist want to interpret and why? This is a questionof individuals, but also of space, time, and . . . More-over, concerning reproduction, we have to admit thatthe copy is never the original, which is why one of us(Hervé This) proposes that the new note by note way ofcooking (see below) should not reproduce, but insteadinvent. It is the same with synthesizers in music: it issure that one can reproduce the sound of a violin, butwhy? If one loves music, why not discover unknownsounds, timbers, and music?

Let’s move to technique now. Here, the project isvery different, because the goal is to make, to build.Techne, in Greek, means to do. For this kind of activ-ity, the criteria are precise reproduction, regularity, andcare. After decades of studying culinary productions,it appeared to one of us that care is the most impor-tant parameter of technicians, because it means givingpleasure, creating a kind of social osmosis through theproduct.

Here, two possibilities exist. One is to reproducewhat was done in the past, by ancestors who lacked ourmodern information on chemistry, physics, and biology;the other is to use all modern information in order to getnew skills and, often, to reach new products.

With odors, these skills can be used in variouscases, such as perfumes, flavors, consumer care prod-ucts, technical preparations, etc. But in all cases, theissue is to deal with mixtures of odorant compounds(i. e., compounds which can link – in one way or an-other – to olfactory receptors) interacting with amatrix,from which components of the mixture are released,at different velocities. For perfumes, this definition isclear, but for flavors? Indeed, if we eat a raspberry,we perceive the flavor of this raspberry. By definition,flavor is the synthetic sensation based on many differ-ent perceptions such as consistency, color, taste, odor,trigeminal perception, and others, because we discovermore and more every year, such as the specific percep-tion of long unsaturated fatty acids on taste buds, or ofcalcium ions, . . . ; and we’re probably not finished.

Odor is one component only, but the status of odor-ant compounds in food is more than odor, because manyfoods have an odor as well as a taste and a trigeminaleffect! It was frequently published in the past that odorwas the most important component of flavor. This is nottrue, as is easily demonstrated by eating after burningone’s mouth with a hot potato, for example: all the fla-vor is lost, even if the odor remains. Let’s keep in mindthat it is not necessary to diminish some (other percep-tions) to make others (odors) appear more important. Ifwe recognize this, we will be able to better include odorin flavor!

Here, let’s conclude by coming back to the idea thatnew information makes new techniques. It is the goal

of this Handbook of Odors to give as fresh informationas possible, and this is why this book is important: newinformation equals modernized techniques.

However, the issue of improving technique is, in-deed, the goal of technology, from techne and logos.One of us published a whole book in order to proposethat there are two kinds of technology, one called lo-cal (the technician, or someone near him, proposes new,more rational ways of doing) and one global, for whichthe engineer looks for results of science, selecting theuseful ones and transferring them to the technical field.Here again, this Handbook is important because by in-cluding chapters from specialists among the best, itgives the basis for innovation. Innovation, the keywordfor the industry!

And finally we look at science. Here we need togive an explanation, because there is much confusionabout it. Indeed, the word science means knowledge,and this is why it is legitimate to speak of the scienceof the shoemaker, or the science of the cook, or the sci-ence of the perfumer . . . However, the science that weneed to discuss here is very different: now, we restrictthe general meaning to the sciences of nature or naturalphilosophy.

For sciences of nature, the goal is to discover themechanisms of phenomena, using the scientific methodbased on:

1. Observing a phenomenon2. Measuring it, which produces a lot of numerical

data3. Grouping these data into laws (i. e., equations)4. Producing theories, i. e., explanations quantitatively

compatible with the data5. Making theoretical predictions which are experi-

mentally tested, and so on, forever.

One could ask if there is a link between science andart, and one of us (Hervé This, certainly a scientist),after decades of close friendship and work with PierreGagnaire (certainly an artist), answers with a verystrong no, even if there has been an intense relationshipbetween artists and scientists, and common characteris-tics such as intuition, liberty, curiosity, enthusiasm . . .Indeed, the goals of science and art are different (mech-anisms on the one hand, emotion on the other), as wellas theways (methodon, in Greek, method) in which theyare achieved.

The method of science was described above; withart it is based on intuition, experience, personal emo-tion, the desire to communicate, . . . If a scientist wantsto move toward art, he or she has to get away from sci-ence to technology, whereas the artist who would liketo move toward science has to go through technique.No relationship between art and science, but rather re-

Foreword VII

lationships between the applications of sciences (verydifferent from the sciences) and the technical compo-nent of art (very different from art, even if it is needed).

All this being said, all of us three agree that onevery important component of our activities is to thinkcorrectly and use good vocabulary, taking the utmostcare with semantics. Very frequently, in building odors,young artists don’t find the precise words, and thishas technical consequences because they cannot selectthe right raw materials. For sciences of nature, also,words are very important, and the great Antoine Laurentde Lavoisier, the creator of modern chemistry, quotedCondillac in order to show why nomenclature was thebasis for advances: sciences explore phenomena, butwe have to think of the phenomena in order to studythem; in order to think we need words, and this is why,Lavoisier said, you cannot improve science without im-proving words and vice versa.

By the way, all of us three also agree that with re-spect to odors or flavors, we would love to have morescientific knowledge.

Imagine that you make an odor without taking intoaccount some trigeminal effect. Imagine that you makefood without pepper: the dish would not be what itcould be. Imagine that you don’t know about someparticular interaction effects of odorants on olfactory re-ceptors: again, the result would not be what it should be.In music, it is as if the piano had only a limited numberof notes. We need more information on the relationshipbetween the molecular structure of odorant compoundsand the odors of such compounds. We need more infor-mation about odor release, which means the physicaland chemical description of the diffusion of odorantsinside and outside matrices, including supramolecularassociations. We need more information on the percep-tion of odorants and the various interactions, beforesmelling them, and on the hedonistic interpretation ofthe olfactory perceptions. Of course, all this holds forfood!

Perfumers succeed empirically with their currentmethods, for example making more vanilla note withbergamot, instead of increasing the quantity of vanillin!This is what two of us (Pierre Kurzenne and MauriceRoucel) name contrast. Perfumers also use metaphoricwords such as hot or sparkling, and in the future it couldbe interesting to better understand what this is.

Concerning all this research, the past makespromises for the future. For odors, Nobel prizes havealready been awarded to advances in the field (terpenes,olfactory receptors), but so much remains to be done!At this point, in particular, the prediction of the odorof a particular chemical structure is out of reach, andthe effect of mixtures of odorants remains elusive. Ofcourse, the exploration of God’s shop is important, but

we have to recognize that nature, in spite of our ideas,is far from perfect! Isn’t plague natural, as are as volca-noes, tsunamis, and so on? If we use clothes, buildings,and perfumes, it is because – in a way – we are fight-ing against nature . . . because human beings belong toculture.

It is strange, in this regard, to classify odors intopoisonous and non-poisonous ones. It is also strangethat some odorant compounds are not as they should be.Why are lead salts sweet, yet toxic? Why are some bit-ternesses accepted (think of beer), yet often associatedwith toxic alkaloids? The same holds for some odor-ants!

Let’s finish with the issue of regulation. It is clearthat the public should be protected –but from what?Let’s first recognize that there is a big difference be-tween danger (a knife is dangerous) and risk (the knifecan be used to kill, but if it is in a closed drawer, thereis no risk). Danger is everywhere, but we have to re-duce risks. In this regard, it is certainly good to knowthe dangers and the exposures in order to make regula-tions – but such rules should focus only on risk, not onhazard.

Now, coming back to the issue of nature/culture,we should say that no product of the perfume or fla-vor industries is natural, because it was produced! Let’sremember that something is natural when it was nottransformed by human beings. If you extract an essencefrom a flower, for example, the flower had to be cul-tivated, and the essence had to be extracted. It’s nolonger natural. But who cares: some products of cultureare much better than natural products. Remember Rem-brandt, Matthias Grünewald, ZaoWo-Ki, Bach, Mozart,Debussy, etc.

Moreover, in this regard, the issue of note by notecooking is probably important for the development ofthe odor industry. This new way of cooking was pro-posed for the first time in 1994 by one of us (HervéThis) and it is the culinary equivalent of synthetic mu-sic. Instead of using flutes and violins, one uses purewaves, in order to make sounds, after which music iscomposed. For cooking, the elementary units are notsound waves, but rather compounds, and note by notecooking does not use traditional ingredients (animal andplant tissues), but rather pure compounds, from whichdishes are made.

This proposal is important for regulation because itkills the need to make particular categories for addi-tives and flavorings: compounds used for making foodwould simply be food ingredients. Where will we findthe needed compounds?Of course they can be extractedor synthesized, and we have to tell this to the public!Extracted or synthesized, vanillin is always vanillin,and water is always water. Of course, today a part of

VIII Foreword

the public fears chemistry, but, indeed, it is becauseit does not understand it. One way to circumvent fearis to make it desirable, trendy, fashionable, even for-bidden. This is how Augustin Parmentier succeededin making the French public eat potatoes, before theFrench Revolution: he invited the king to eat them!This strategy has been used to implement note by notecooking.

Let’s imagine that we succeed in developing thisnew culinary trend. How shall we cook? A good waybeing explored today is to design the shapes and con-sistencies, then to design color, taste, odor and trigemi-nal perception, as we add spices to traditional dishes.However flavorists know too well that knowledge isneeded in order to get the desired sensation, becauseof the chemical and physical interactions with the var-ious compartments of the matrix. Chefs will have tolearn, which means that flavorists’ collaboration will beneeded. Of course, one could imagine using flavoringsas we use aromatic herbs and spices, but wouldn’t it bemuch more interesting to use pure solutions of partic-ular odorants (for example, a very small concentration

of 1-cis-hexen-3-ol in ordinary oil), or kits of odors, inorder to make entirely new flavors (edible perfumes)?By experience, chefs are not ready to have flavoriststake the lead on their productions, which means that thefirst, traditional solution of using flavorings is probablynot the future. If kits or pure solutions are the future,the odor industry has to be ready to make entirely newproducts.

Finally, after this long tour, we have to recognizethat life is wonderful, in particular because the worldof odors is fascinating. About art, for technique, fortechnology, for science, there are so many open ques-tions! Questions are promises of answers – if we workwith passion! Passion for odors, passion for imagina-tion, passion for emotions, passion for knowledge . . .

In mathematics, the great David Hilbert said Wirmüssen wissen, wir werden wissen (we must know, wewill know)!

Pierre KurzenneMaurice RoucelHervé This

IX

Preface

Olfaction in humans is a sensory dimension that is of-ten underestimated or overlooked entirely. The senseof smell is commonly believed to play only a mi-nor role in human perceptual experiences, which isan understandable misconception when considering theeveryday dominance of vision and hearing for commu-nication, augmented by touch for interaction with theimmediate surroundings. Moreover, these more promi-nent sensory modalities are known to exert both physi-ological and psychological influences on humans (e.g.,sound/noise, light/vision, and temperature/climate) thatcan have an impact on our general wellbeing. In ad-dition, our sense of smell is typically considered tobe inferior to olfaction found elsewhere in the ani-mal kingdom; for example, numerous species are ableto trace the path of an odor source over great dis-tances, or habitually undergo behavioral responses thatare strongly modulated by volatile chemicals, e.g.,pheromone-type compounds, that may play a role inaggression, mating, or rearing offspring. An absenceof scientific proof does not preclude such remarkablecapabilities in humans, yet there is widespread dis-regard for the importance and impact of the humansense of smell. Seemingly, this most likely relates toits association with primitive, animal-like behavior andpotentially uncontrollable effects and responses that areelicited by smells, which are considered to be less ra-tional and, therefore, more appropriate for the animalkingdom; surely the higher intellect of homo sapiens isnot prone to behavioral responses from such primitiveinfluences!

Yet, there is more to the human sense of smellthan meets the nose, so to speak. There is an everincreasing body of evidence from ground-breaking re-search discoveries that demonstrates the crucial rolethat smells play in shaping our lives. From birth on-wards we learn to interact with our environment usingour sense of smell. Evolutionary processes have engen-dered a multifaceted communication that is supported– even dominated – by olfaction. This might be in theform of the smells that nurture the relationship betweenmother and child or influence partner selection, aromasthat form our food preferences, or odors that warn us ofdangers.

In the modern world we are increasingly exposedto smells that were not encountered by our ancestors.These are ubiquitous in our present-day environments

and are met in all aspects of daily life, with sourcesranging from manmade materials, industry, transport,household products, etc.; the list is practically endless.The outcome of this constant evolutionary process inmaterial, product and application development is thatwe have generated a tolerance or even unawareness tonumerous modern smells despite their often pervasiveand abundant nature. By contrast, smells that relate toour appreciation of products such as foods have at-tracted intense scientific interest spanning decades, withthe earliest discoveries dating back to when chemistrywas still in its infancy. Specifically, pleasantly-smellingraw materials and compounds considered to be attrac-tive to humans (e.g., as body scents or room fragrances)were at the focus of early research, sometimes withenormous efforts made to recover and enrich substancesto a sufficient extent to enable a chemo-analytical elu-cidation of the underlying odorous molecules; suchefforts were especially laborious and time-consumingat a time when the analytics and respective instrumen-tation were still rudimentary. Nowadays, an array ofmethods are at our disposal to resolve even the mostcomplex of odor mixtures and decode the structures ofindividual molecules at extremely low concentrations;thus, rather than the insufficient sensitivities and res-olutions limiting early research in this field, presentodorant analytics might be considered as searching fora needle in a haystack, with individual odorants presentamongst a forest of competing signals.

With emerging progress in the biochemical,biomedical, and neurosciences, research in olfactionhas subsequently expanded to include a strong focuson the impact of smells on humans. This new direc-tion has revealed important insights into how smellsare perceived, processed and memorized, and how odorimpressions influence our everyday lives. Nevertheless,the nature of numerous odorants remains unknown, asdoes their influence on perception, physiology and well-being. This is especially true for the modern smells thatare encountered on a daily basis at home, at work, orwhen out and about.

A comprehensive treatment of smells – from an-cient to modern and rare to common – in relation to theimpact they have on our lives currently does not exist.This handbook aims to bridge this gap by aligning thesenso-chemo-analytical characterization of smells en-countered by mankind, tracing the diverse routes of po-

X Preface

tential formation and release pathways, and elucidatingthe perceptual hedonic, behavioral, and physiologicalresponses of humans to such odors at different stagesof life. This book is intended to build a foundation fora hitherto widely overlooked area of research that haswider ramifications for human life, and to instigate in-

tensified interdisciplinary discussions as a catalyst forgaining further insights and discoveries.

Andrea BuettnerMunich, GermanyOctober 2016

XI

About the Editor

Andrea Buettner read for an undergraduate degree in food chemistry at LudwigMaximilian University of Munich, Germany, and then completed her postgraduate andpostdoctoral research at both the German Research Center for Food Chemistry (DFA)and the Technical University of Munich from 1995 to 2002. Following her habilitationto qualify for full professorship in 2007 she was appointed and still holds two concur-rent positions, both as Founder and Head of the Department of Sensory Analytics atthe Fraunhofer Institute for Process Engineering and Packaging IVV in Freising, Ger-many, and of the Odor and Aroma Research Group at Friedrich-Alexander-Universität(FAU) Erlangen-Nürnberg, Germany. Andrea has received several accolades for herresearch, including the Kurt-Täufel Prize for young scientists from the Food Chem-istry Division of the German Chemistry Society (2010), the Young Investigator Awardfrom the Food and Agricultural Division of the American Chemical Society (2011),the Danone Innovation Prize (together with Caroline Siefarth, 2012), and the NutriciaScience Award (2013). Since 2012, Andrea has held a position as Full Professor ofAroma Research at FAU in Erlangen.

Andrea’s expertise encompasses the characterization of the main odor triggers in aromas, flavors, and commonodors. Specifically, Andrea is renowned for identifying and characterizing the primary odor triggers in typicalaromas, flavors, fragrances, and smells that are associated with everyday life. Amongst other things, her workinvolves disentangling and subsequently reconstructing characteristic smells, from food aromas to modern man-made materials, based on combined chemo-analytical and human-sensory characterization of the molecular odorconstituents. Her specialized field extends to characterizing odorant release via on-line monitoring of distributionand delivery processes. This particular aspect of her work, for example, has provided insights into the impor-tance of the combined effects of the food matrix, saliva, mucosa, mastication, and swallowing on flavor releaseand perception. In view of physiological processes related to odorant exposure and uptake, recent investigationshave targeted pharmacokinetic aspects of odorant inhalation, absorption, biotransformation, and elimination viabreath, urine, sweat, and human milk. Accordingly, Andrea’s research aims to raise awareness of smell and odordimensions in human life, especially in common, everyday situations. Her goal is to cross-link technological,chemical, or physiological aspects and to provide novel solutions in odor research. Novel or modified techniques,methodologies, processing technologies, analytical tools, and approaches related to smell are further outcomes ofher research strategy, with the ultimate aim to nourish interdisciplinary cross-talk on a topic that links chemical,analytical, material, and process engineering sciences with the fields of sociology and socio-ecology, psychology,and physiology.

XIII

List of Authors

Artin ArshamianKarolinska InstitutetDepartment of Clinical NeuroscienceNobels Väg 917177 Stockholm, Swedenand

Radboud UniversityCentre for Language Studies, and DondersInstitute for Brain, Cognition, and Behaviour6500 Nijmegen, The Netherlandsartin.arshamian@ki.se

Anat ArziThe Weizmann Institute of ScienceDepartment of Neurobiology234 Herzl St.76100 Rehovot, Israelanat.arzi@weizmann.ac.il

Jinhe BaiU.S. Dept. of Agriculture – Agricultural ResearchServiceUS Horticultural Research Laboratory2001 South Rock RoadFort Pierce, FL 34945, USAjinhe.bai@ars.usda.gov

Nicolas BaldoviniUniversité de Nice-Sophia AntipolisInstitut de Chimie de Nice, UMR 7272 CNRS28 Avenue Valrose06108 Nice, Francebaldovin@unice.fr

Elisabeth BaldwinU.S. Dept. of Agriculture – Agricultural ResearchServiceUS Horticultural Research Laboratory2001 South Rock RoadFort Pierce, FL 34945, USAliz.baldwin@ars.usda.gov

Jonathan BeauchampFraunhofer Institute for Process Engineering andPackaging (IVV)Giggenhauser Strasse 3585354 Freising, Germanyjonathan.beauchamp@ivv.fraunhofer.de

Ofir BenjaminTel Hai CollegeFood Science Department1220800 Upper Galilee, Israelofirbe@telhai.ac.il

Carlo BicchiUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italycarlo.bicchi@unito.it

Roland BlachFraunhofer Institute for Industrial Engineering IAONobelstrasse 1270569 Stuttgart, Germanyroland.blach@iao.fraunhofer.de

Sanne BoesveldtWageningen UniversityDivision of Human NutritionStippeneng 46708 WE, Wageningen, The Netherlandssanne.boesveldt@wur.nl

Heinz BreerUniversity of HohenheimInstitute of PhysiologyGarbenstrasse 3070593 Stuttgart, Germanybreer@uni-hohenheim.de

Andrea Burdack-FreitagFraunhofer Institute for Building PhysicsDepartment Building Chemistry, Building Biology,HygieneFraunhoferstrasse 1083626 Valley, Germanyandrea.burdack-freitag@ibp.fraunhofer.de

Cecilia CaglieroUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italycecilia.cagliero@unito.it

XIV List of Authors

Annachiara CavazzanaMonell Chemical Senses Center3500 Market StreetPhiladelphia, PA 19104, USAacavazzana@monell.org

Cinzia CecchettoMonell Chemical Senses Center3500 Market StreetPhiladelphia, PA 19104, USAccecchet@sissa.it

Pierre ChatelainChemCom SARoute de Lennik 8021070 Brussels, Belgiumpch@chemcom.be

Sung-Tong ChinImperial College LondonDepartment of Surgery and CancerNorfolk PlaceLondon, W2 1PG, UKs.chin@imperial.ac.uk

Norbert ChristophBavarian Health and Food Safety Authority (LGL)Luitpoldstrasse 197082 Würzburg, Germanynorbert.christoph@lgl.bayern.de

Géraldine CoppinYale UniversityThe John B. Pierce Laboratory290 Congress AvenueNew Haven, CT 06519, USAgeraldine.coppin@yale.edu

Chiara CorderoUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italychiara.cordero@unito.it

Michael CzernyFrauenhofer Institute for Process Engineering andPackagingGiggenhauser Strasse 3585354 Freising, Germanymichael.czerny@ivv.fraunhofer.de

Manfred DangelmaierFraunhofer Institute for Industrial Engineering IAONobelstrasse 1270569 Stuttgart, Germanymanfred.dangelmaier@iao.fraunhofer.de

Philippe DarrietBordeaux UniversityInstitute of Vine and Wine Sciences210 chemin de Leysott, CS 5000833882 Villenave d’Ornon, Francephilippe.darriet@u-bordeaux.fr

Sylvain DelplanqueUniversity of GenevaSwiss Center for Affective Sciences9 Chemin des Mines1202 Geneva, Switzerlandsylvain.delplanque@unige.ch

Richard L. DotyUniversity of PennsylvaniaPerelman School of Medicine5 Ravdin PavilionPhiladelphia, PA 19104, USAdoty@mail.med.upenn.edu

Michael DreselAB InBev Belgium n.v.Brouwerijplein 1 – GITeC3000 Leuven, Belgiummichael.dresel@hotmail.de

Graham EllisGivaudan International SARegulatory Affairs and Product Safety5 Rue de la Parfumerie1214 Vernier, Switzerlandgraham.ellis@givaudan.com

Karl-Heinz EngelTechnical University of MunichChair of General Food TechnologyMaximus-von-Imhof-Forum 285354 Freising-Weihenstephan, Germanyk.h.engel@wzw.tum.de

Graham T. EyresUniversity of OtagoDepartment of Food Science9054 Dunedin, New Zealandgraham.eyres@otago.ac.nz

List of Authors XV

Jitka FialováCharles UniversityFaculty of ScienceViničná 712844 Prague, Czech Republicand

National Institute of Mental Health25067 Klecany, Czech Republicjitka.fialova@nudz.cz

Jean-Jacques FilippiUniversité de Nice-Sophia AntipolisInstitut de Chimie de Nice, UMR 7272 CNRS28 Avenue Valrose06108 Nice, Francejfilippi@unice.fr

Jörg FleischerUniversity of HohenheimInstitute of PhysiologyGarbenstrasse 3070593 Stuttgart, Germanyjoergf@uni-hohenheim.de

Johannes FrasnelliUniversité du Québec à Trois-RivièresDepartment of Anatomy3351 boul. des ForgesTrois-Rivières, QC G9A 5H7, Canadajohannes.a.frasnelli@uqtr.ca

Jessica FreiherrRWTH Aachen UniversityDiagnostic and Interventional NeutoradiologyPauwelstrasse 3052074 Aachen, Germanyjfreiherr@ukaachen.de

Eric FrérotFirmenich SAAnalytical Innovation Corp. R&D1, Route Des JeunesCH-1211, Geneva, Switzerlanderic.frerot@firmenich.com

Kristina FriedlandFriedrich-Alexander-UniversitätErlangen-NürnbergDepartment of Chemistry and PharmacyCauerstrasse 691058 Erlangen, Germanykristina.leuner@fau.de

Marcus A. GlombMartin-Luther University Halle-WittenbergInstitute of ChemistryKurt-Mothes-Str. 206120 Halle, Germanymarcus.glomb@chemie.uni-halle.de

Amy R. GordonKarolinska InstitutetDepartment of Clinical NeuroscienceNobels Väg 917177 Stockholm, Swedenamy.gordon@ki.se

Brian GuthrieCargill Inc.Global Food Reserach2301 Crosby RoadWayzata, MN 55391, USAbrian_guthrie@cargill.com

Ute HabelUniklinik RWTH AachenDepartment of Psychiatry, Psychotherapy andPsychosomaticsPauwelsstrasse 3052074 Aachen, Germanyuhabel@ukaachen.de

Ilana S. HairstonAcademic College of Tel Aviv YaffoDepartment of Behavioral Sciences2 Rabenu Yerucham St.Tel Aviv, Israelilanahai@mta.ac.il

Christian Harteneck (deceased)

Hanns HattRuhr-University BochumDepartment of Cell PhysiologyUniversitätsstrasse 15044801 Bochum, Germanyhanns.hatt@ruhr-uni-bochum.de

Heike HauschildtOlfasense GmbHFraunhofer Strasse 1324118 Kiel, Germanyhhauschildt@olfasense.com

XVI List of Authors

Jan HavlíčekCharles UniversityFaculty of ScienceViničná 712844 Prague, Czech Republicjhavlicek@natur.cuni.cz

Anja HeinleinFriedrich-Alexander-UniversitätErlangen-NürnbergDepartment of Chemistry and PharmacyHenkestrasse 991054 Erlangen, Germanyanja.heinlein@fau.de

Thomas HummelTU DresdenSmell & Taste Clinic, Department ofOtorhinolaryngologyFetscherstrasse 7401307 Dresden, Germanythummel@mail.zih.tu-dresden.de

Cornell KärnekullStockholm UniversityDepartment of PsychologyFrescati Hagväg 9A10691 Stockholm, Swedenstina.cornell.karnekull@psychology.su.se

Gerhard KrammerSymrise AGMühlenfeldstrasse 137603 Holzminden, Germanygerhard.krammer@symrise.com

Basile N. LandisUniverstiy of Geneva Medical School and GenevaUniversity HospitalsDepartment of OtorhinolaryngologyRue Gabrielle-Perret-Gentil 41205 Geneva, Switzerlandbasile.landis@hcuge.ch

Maria LarssonStockholm UniversityDepartment of PsychologyFrescati Hagväg 9A10691 Stockholm, Swedenmarlar@psychology.su.se

Matthias LaskaLinköping UniversityIFM BiologyCampus Valla58183 Linköping, Swedenmalas@ifm.liu.se

Erica LibertoUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italyerica.liberto@unito.it

Johan N. LundströmKarolinska InstitutetDepartment of Clinical NeuroscienceNobels Väg 917177 Stockholm, Swedenjohan.lundstrom@ki.se

Simona ManescuUniversité de MontréalDepartment of Psychology90 avenue Vincent d’IndyMontréal, QC H2V 2S9, Canadasimona.manuescu@umontreal.ca

Bettina MannebeckOlfasense GmbHFraunhofer Strasse 1324118 Kiel, Germanybmannebeck@olfasense.com

Dietmar MannebeckOlfasense GmbHFraunhofer Strasse 1324118 Kiel, Germanydmannebeck@olfasense.com

Philip J. MarriottMonash UniversityAustralian Centre for Research on SeparationScienceWellington RoadVictoria, 3800, Australiaphilip.marriott@monash.edu

Florian MayerFraunhofer Institute for Building PhysicsDepartment Building Chemistry, Building Biology,HygieneFraunhoferstrasse 1083626 Valley, Germanyflorian.mayer@ibp.fraunhofer.de

List of Authors XVII

Andreas NatschGivaudan Schweiz AGBiosciencesUeberlandstrasse 1388600 Duebendorf, Switzerlandandreas.natsch@givaudan.com

Nathalie NibbeOlfasense GmbHFraunhoferstrasse 1324118 Kiel, Germanynnibbe@olfasense.com

Johannes NieblerFriedrich-Alexander-UniversitätErlangen-NürnbergDepartment of Chemistry and PharmacyHenkestrasse 991054 Erlangen, Germanyjohannes.niebler@fau.de

Jeannette Nuessli GuthETH ZurichDept. of Health Sciences and TechnologySchmelzbergstrasse 98092 Zurich, Switzerlandjnuessli@ethz.ch

Mats J. OlssonKarolinska InstitutetDepartment of Clinical NeuroscienceNobels Väg 917177 Stockholm, Swedenmats.j.olsson@ki.se

Ulrich R. OrthKiel UniversityA&F Marketing – Consumer PsychologyWilhelm-Seeling-Platz 724098 Kiel, Germanyuorth@ae.uni-kiel.de

Jane K. ParkerUniversity of ReadingThe Flavour CenterWhiteknights CampusReading, RG6 6AP, UKj.k.parker@rdg.ac.uk

Valentina ParmaMonell Chemical Senses Center3500 Market StreetPhiladelphia, PA 19104, USAvparma@monell.org

Bettina M. PauseUniversity of DüsseldorfDepartment of Experimental PsychologyUniversitätsstrasse 140225 Düsseldorf, Germanybettina.pause@hhu.de

Ofer PerlThe Weizmann Institute of ScienceDepartment of Neurobiology234 Herzl St.76100 Rehovot, Israelofer.perl@weizmann.ac.il

Magali PhilippeauChemCom SARoute de Lennik 8021070 Brussels, Belgiummph@chemcom.be

Wilhelm PickenhagenCh. Montmusy 3CH 1290, Chavannes-des-Bois, Switzerlandpickenhagen@bluewin.ch

Anne PlottoU.S. Dept. of Agriculture – Agricultural ResearchServiceUS Horticultural Research Laboratory2001 South Rock RoadFort Pierce, FL 34945, USAanne.plotto@ars.usda.gov

Alexandre PonsBordeaux UniversityInstitute of Vine and Wine Sciences210 chemin de Leysott, CS 5000833882 Villenave d’Ornon, Franceand

Seguin Moreau Cooperage16103 Cognac, Francealexandre.pons@u-bordeaux.fr

Christina RegenbogenUniklinik RWTH AachenDepartment of Psychiatry, Psychotherapy andPsychosomaticsPauwelsstrasse 3052074 Aachen, Germanycregenbogen@ukaachen.de

XVIII List of Authors

Gary ReinecciusUniversity of MinnesotaDepartment of Food Science and Nutrition1334 Eckles Ave.Saint Paul, MN 55108, USAgreinecc@umn.edu

Martin RichterFraunhofer Research Institution for Modular SolidState Technologies EMFTHansastrasse 27d80686 Munich, Germanymartin.richter@emft.fraunhofer.de

S. Craig RobertsUniversity of StirlingDivision of PsychologyStirling, FK9 4LA, UKcraig.roberts@stir.ac.uk

Philippe RombauxUniversity of Louvain – BrusselsDepartment of Otorhinolaryngology, CliniquesUniversitaires Saint LucAvenue Hippocrate 101200 Woluwe-Saint-Lambert, Belgiumphilippe.rombaux@uclouvain.be

Patrizia RubioloUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italypatrizia.rubiolo@unito.it

Maren RunteZHAW Zurich University of Applied SciencesSchool of Applied LinguisticsTheaterstrasse 15c8400 Winterthur, Switzerlandmaren.runte@zhaw.ch

Michael RychlikTechnical University of MunichZIEL Research Center for Nutrition and FoodSciencesAlte Akademie 1085354 Freising, Germanymichael.rychlik@tum.de

Christian SallesUniversité Bourgogne Franche-Comté (UBFC)Centre for Taste and Feeding Behavior17 Rue Sully21000 Dijon, Francechristian.salles@dijon.inra.fr

David SanderUniversity of GenevaSwiss Center for Affective Sciences9 Chemin des Mines1202 Geneva, Switzerlanddavid.sander@unige.ch

Elise Sarrazin50 Rue des Carrières92150 Surenes, Franceelise.sarrazin.marechal@gmail.com

Benoist SchaalUniversity of BurgundyCentre of Taste and Feeding Behaviour9E Boulevard Jeanne d’Arc21000 Dijon, Francebenoist.schaal@u-bourgogne.fr

Antje SchellenbergBavarian Health and Food Safety Authority (LGL)Veterinärstrasse 285674 Oberschleissheim, Germanyantje.schellenberg@lgl.bayern.de

Boris SchillingGivaudan Schweiz AGFragrance S&T – BiosciencesÜberlandstrasse 138Duebendorf, Switzerlandboris.schilling@givaudan.com

Han-Seok SeoUniversity of ArkansasDept. of Food Science2650 North Young AvenueFayetteville, AR 72704, USAhanseok@uark.edu

Janina SeubertKarolinska InstitutetDepartment of Clinical NeuroscienceNobels väg 917177 Stockholm, Swedenjanina.seubert@ki.se

Anne-Marie SeuvreUniversity of Burgundy/Agrosup DijonUMR PAM Food Processing and Microbiology1 Esplanade Erasme21000 Dijon, Franceanne-marie.seuvre@u-bourgogne.fr

List of Authors XIX

Barbara SgorbiniUniversity of TorinoDept. of Drug Science and TechnologyVia Pietro Giuria 910125 Torino, Italybarbara.sgorbini@unito.it

Noam SobelThe Weizmann Institute of ScienceDepartment of Neurobiology234 Herzl St.76100 Rehovot, Israelnoam.sobel@weizmann.ac.il

Veronika SomozaUniversity of ViennaDepartment of Nutritional and PhysiologicalChemistryAlthanstrasse 141090 Wien, Austriaveronika.somoza@univie.ac.at

Marc SpehrRWTH Aachen UniversityDept. of ChemosensationWorringerweg 352074 Aachen, Germanym.spehr@sensorik.rwth-aachen.de

Christian StarkenmannFirmenich SARoute des Jeunes 11211 Geneva, Switzerlandchristian.starkenmann@firmenich.com

Jörg StrotmannUniversity of HohenheimInstitute of PhysiologyGarbenstrasse 3070593 Stuttgart, Germanystrotman@uni-hohenheim.de

Katinka TemmeUniversity of Applied Sciences AugsburgFaculty of Architecture and Civil EngineeringAn der Hochschule 186161 Augsburg, Germanykatinka.temme@hs-augsburg.de

Michael ThielSana-Klinikum Remscheid GmbHAcademical Teaching Hospital of the University ofCologneBurger Strasse 21142859 Remscheid, Germanymichael.thiel@sana.de

Alex VeithenChemCom SARoute de Lennik 8021070 Brussels, Belgiumave@chemcom.be

Sophie VeitingerRuhr-University BochumDepartment of Cell PhysiologyUniversitätsstrasse 15044801 Bochum, Germanyherlisb6@rub.de

Andrée VoilleyUniversity of Burgundy/Agrosup DijonUMR PAM Food Processing and Microbiology1 Esplanade Erasme21000 Dijon, Franceandree.voilley@u-bourgogne.fr

Jessica WalkerUniversity of ViennaDepartment of Nutritional and PhysiologicalChemistryAlthanstrasse 141090 Wien, Austriajessica.walker@univie.ac.at

Donald WilsonNYU Scool of MedicineDepartment of Child and Adolescent Psychiatry1 Park AvenueNew York, NY 10016, USAdonald.wilson@nyumc.org

Matthias WüstUniversity of BonnInstitute of Nutritional and Food SciencesEndenicher Allee 11-1353115 Bonn, Germanymatthias.wuest@uni-bonn.de

Chahan YeretzianZHAW Zurich University of Applied SciencesInstitute of Chemistry and BiotechnologyEinsiedlerstrasse 31CH-8820, Wädenswil, Switzerlandyere@zhaw.ch

Wiebke ZanderCargill GmbHSeehafenstrasse 221079 Hamburg, Germanywiebke_zander@cargill.com

XX List of Authors

Erika ZardinFraunhofer Institute for Process Engineering andPackaging (IVV)Giggenhauser Strasse 3585354 Freising, Germanyerika.zardin@ivv.fraunhofer.de

XXI

Contents

List of Abbreviations ............................................................. XXXIII

1 History of Odor and OdorantsWilhelm Pickenhagen ........................................................... 11.1 Defining Odor and Odorants............................................ 11.2 The Chemical Senses .................................................... 11.3 History of Use of Odorants .............................................. 2References ....................................................................... 8

Part A Molecular Aspects and Formation Pathways

2 Biosynthesis of Plant-Derived OdorantsMatthias Wüst ................................................................... 132.1 Biosynthesis of Plant-Derived Odorants ............................... 132.2 Constitutive Biosynthetic Pathways .................................... 162.3 Stress-Induced Biosynthesis of Plant Volatiles ....................... 332.4 Outlook................................................................... 33References ....................................................................... 33

3 Natural Fragrant Raw MaterialsNicolas Baldovini, Jean-Jacques Filippi ....................................... 393.1 Identification of Odor Active Constituents in Natural Raw Materials 403.2 Odor-Active Constituents of Selected Natural Raw Materials ........ 443.3 Conclusion ............................................................... 57References ....................................................................... 57

4 Incense MaterialsJohannes Niebler ................................................................ 634.1 Selected Incense Materials ............................................. 654.2 Incense Preparations.................................................... 794.3 Benefits and Hazards of Incense Use .................................. 814.4 Conclusion and Outlook ................................................ 83References ....................................................................... 83

5 Mechanistic Pathways of Non-Enzymatic Flavor FormationMarcus A. Glomb ................................................................ 875.1 Maillard Reaction – General Considerations .......................... 875.2 ˛-Dicarbonyl Compounds .............................................. 895.3 Strecker Degradation .................................................... 935.4 Other Mechanisms....................................................... 995.5 Conclusions .............................................................. 102References ....................................................................... 103

XXII Contents

Part B Food and Flavors

6 CoffeeChahan Yeretzian ................................................................ 1076.1 Coffee Aroma – From the Seed to the Cup ............................ 1076.2 The Sensory Experience of Coffee ...................................... 1086.3 Coffee Aroma Compounds .............................................. 1106.4 Analytical Techniques for Coffee Aroma Analysis ..................... 1156.5 Trends and New Developments in Coffee Aroma Analysis............ 1166.6 What Next?............................................................... 122References ....................................................................... 122

7 BeerMichael Dresel ................................................................... 1297.1 Raw Materials............................................................ 1307.2 Flavor Evolution of Hoppy Aroma ...................................... 1327.3 Special Flavors ........................................................... 1357.4 Influence on the Sensory Sensation due to Other Constituents ..... 1397.5 Outlook................................................................... 139References ....................................................................... 139

8 WinePhilippe Darriet, Alexandre Pons............................................... 1438.1 Composition of the Wine Matrix ....................................... 1448.2 Perceptual Interaction Phenomena.................................... 160References ....................................................................... 162

9 FruitsAnne Plotto, Jinhe Bai, Elisabeth Baldwin .................................... 1719.1 Fruit Volatiles ............................................................ 1719.2 Impact of Volatiles on Fruit Flavor ..................................... 1759.3 Flavor of Specific Fruits ................................................. 1779.4 Conclusion ............................................................... 183References ....................................................................... 183

10 MeatJane K. Parker ................................................................... 19110.1 Introduction to Meat Flavor ............................................ 19110.2 Characterization of Meat Aroma........................................ 19210.3 Mechanisms of Aroma Generation ..................................... 20110.4 Factors Affecting Meat Flavor ........................................... 210References ....................................................................... 215

11 Fats and OilsEric Frérot ........................................................................ 22311.1 Analytical Techniques Used to Study Aroma Compounds

in Fats and Oils .......................................................... 22411.2 The Different Chemical Classes of Aroma Compounds

Found in Fats and Oils, and How They Are Formed .................. 23011.3 The Aroma of Fats and Oils ............................................. 23811.4 Conclusion ............................................................... 256References ....................................................................... 256

Contents XXIII

12 Aroma Encapsulation and Controlled DeliveryGary Reineccius .................................................................. 26112.1 Diversity in Aroma Compounds......................................... 26212.2 Chemical Reactions that Lead to Flavor Loss .......................... 26212.3 Evaporation as a Mechanism of Flavor Loss ........................... 26412.4 Techniques for Preserving Flavor....................................... 26412.5 Controlled Delivery of Food Aroma..................................... 26712.6 Unmet Needs ............................................................ 26912.7 Conclusions .............................................................. 270References ....................................................................... 270

13 Physico-Chemical Interactions in the Flavor-Release ProcessAnne-Marie Seuvre, Andrée Voilley ............................................ 27313.1 Nature of the Interactions Aroma Compounds-Matrices ............. 27413.2 Physico-Chemical Characteristics of Aroma Compounds ............. 27713.3 Composition and Properties of the Matrices .......................... 28013.4 Methods to Study Interactions and Their Role on Transfers.......... 28813.5 Flavor Release or Retention ............................................ 29213.6 Preservation of Food Quality and Perspectives ....................... 294References ....................................................................... 295

14 Models of the Oral Cavity for the Investigation of OlfactionChristian Salles, Ofir Benjamin ................................................. 30314.1 Oral Food Processing and the Effect on Olfaction ..................... 30314.2 Simulation of Oral Processing .......................................... 30614.3 Conclusions .............................................................. 315References ....................................................................... 316

15 Regulatory Oversight and Safety Assessment of FlavoringsKarl-Heinz Engel ................................................................ 31915.1 Regulatory Framework .................................................. 31915.2 Safety Assessment Procedures.......................................... 32215.3 Flavorings Other than Flavoring Substances .......................... 32615.4 Outlook................................................................... 326References ....................................................................... 326

16 Odors in Paper and Cardboard PackagingMichael Czerny ................................................................... 32916.1 Paper ..................................................................... 32916.2 Legal Basics: Food Legislation .......................................... 33116.3 Analytical Methods ...................................................... 33216.4 Off-Odorants in Paper and Cardboard Packaging ..................... 33716.5 Summary ................................................................. 339References ....................................................................... 339

Part C Analytics, Sensor Technologyand Human-Sensory Evaluation

17 Gas Chromatography-Mass Spectrometry in Odorant AnalysisSung-Tong Chin, Graham T. Eyres, Philip J. Marriott ......................... 34317.1 Rationale for Multidimensional GC .................................... 344

XXIV Contents

17.2 Offline MDGC-O .......................................................... 34417.3 Online MDGC-O .......................................................... 34617.4 Novel Approaches in Olfactometry ..................................... 34717.5 GC � GC Approaches for Global Volatile Screening .................... 34817.6 Integrated MDGC-O Arrangement ...................................... 34917.7 Current Challenges and Future Perspectives .......................... 35017.8 Concluding Remarks .................................................... 350References ....................................................................... 350

18 Odorant Detection by On-line Chemical IonizationMass SpectrometryJonathan Beauchamp, Erika Zardin ........................................... 35518.1 Techniques ............................................................... 35618.2 Applications.............................................................. 36918.3 Conclusion and Outlook ................................................ 391References ....................................................................... 393

19 Enantioselective Gas Chromatographywith Cyclodextrin in Odorant AnalysisCecilia Cagliero, Barbara Sgorbini, Chiara Cordero, Erica Liberto,Patrizia Rubiolo, Carlo Bicchi................................................... 40919.1 Chiral Recognition and Enantioselective

Gas-Chromatography (Es-GC)........................................... 41119.2 Measurement of the Enantiomeric Distribution ...................... 41719.3 Enantioselective GC Analysis with Cyclodextrins

in the Flavor and Fragrance Field ...................................... 41819.4 Testing Column Efficiency and Enantioselectivity..................... 41819.5 Analysis of Enantiomers in Complex Samples......................... 42019.6 Fast Enantioselective GC Analysis ...................................... 42419.7 Total Analysis Systems and Real-World Sample Analysis............. 43219.8 Conclusions .............................................................. 433References ....................................................................... 433

20 Stable Isotope Ratio Analysis for Authenticity ControlNorbert Christoph, Antje Schellenberg, Wiebke Zander, Gerhard Krammer . 43920.1 Fundamentals ........................................................... 44020.2 Methods for the Determination of Isotope Ratios .................... 44420.3 Selected Applications of Authentication of Odorants................. 44620.4 Requirements and Guidelines for Authentication .................... 45220.5 Conclusion ............................................................... 454References ....................................................................... 454

21 Machine OlfactionBrian Guthrie .................................................................... 45921.1 Chemical Senses ......................................................... 46021.2 Sensor Types ............................................................. 46321.3 Biomimetic Data Analytic Approaches ................................. 47321.4 Machine Olfaction Applications ........................................ 47421.5 Conclusions .............................................................. 482References ....................................................................... 482

Contents XXV

22 High-Throughput Receptor Screening AssayAlex Veithen, Magali Philippeau, Pierre Chatelain............................ 50522.1 Working with ORs........................................................ 50522.2 How to Identify Ligands with in vitro Functional Assays............. 50722.3 Applications.............................................................. 51422.4 Other Chemosensory Receptors ........................................ 51922.5 Concluding Remarks .................................................... 521References ....................................................................... 522

23 Psychophysical Testing of Human Olfactory FunctionRichard L. Doty .................................................................. 52723.1 Stimulus Presentation Procedures ..................................... 52723.2 Measurement of Basal Olfactory Sensitivity ........................... 52823.3 Measurement of Suprathreshold Odor Perception .................... 53123.4 Issues in Olfactory Psychophysical Measurement ..................... 53723.5 Concluding Remarks .................................................... 539References ....................................................................... 539

24 Olfactometers According to EN 13725Dietmar Mannebeck ............................................................. 54524.1 General Characteristics .................................................. 54624.2 Evaluation and Presentation of Measurement Results ............... 551References ....................................................................... 551

25 Assessment of Environmental Odor ImpactsBettina Mannebeck, Heike Hauschildt ......................................... 55325.1 Odor Measurement at the Source ...................................... 55425.2 Measurement of Odor Impact by Field Measurement ................ 55725.3 Examples of the Evaluation of Odor and Emissions in Ambient Air . 56025.4 Conclusion ............................................................... 561References ....................................................................... 561

26 Material Odor Emissions and Indoor Air QualityAndrea Burdack-Freitag, Anja Heinlein, Florian Mayer ...................... 56326.1 Effects on Indoor Air Quality............................................ 56426.2 Odor Evaluation Methods .............................................. 56426.3 Indoor Odorants – Selected Sources ................................... 56926.4 Odor Avoidance and Reduction Measures ............................. 57526.5 Health Aspects and Sensory Irritations ................................ 576References ....................................................................... 577

Part D Odorant Sensing and Physiological Effects

27 Odorant SensingHeinz Breer, Jörg Fleischer, Jörg Strotmann ................................... 58527.1 Structure of the Olfactory System ...................................... 58527.2 Olfactory Sensory Neurons .............................................. 58627.3 Recognition of Odorants ................................................ 58827.4 Expression of Odorant Receptor Genes ................................ 59227.5 Odorant Receptor-Specific Wiring of Olfactory Sensory Neurons .... 595References ....................................................................... 598

XXVI Contents

28 Nasal Periceptor ProcessesBoris Schilling ................................................................... 60528.1 Xenobiotic-Metabolizing Enzymes in the Olfactory Epithelium ..... 60828.2 Cytochrome P450 Enzymes ............................................. 60928.3 Exploring the Substrate and Inhibitor Range

of Olfactory P450 Enzymes ............................................. 60928.4 Evidence for the Role of Biotransformation Enzymes

in Olfaction from Animal Studies ...................................... 61128.5 Human Sensory Studies ................................................. 61228.6 Discussion................................................................ 613References ....................................................................... 614

29 Metabolism of Odorants in HumansMichael Rychlik .................................................................. 61729.1 General Principles of Metabolism and Absorption ................... 61829.2 Metabolism of Important Classes of Odorants ........................ 61929.3 Conclusions and Outlook for Further Research........................ 626References ....................................................................... 627

30 Olfactory SubsystemsMarc Spehr ....................................................................... 63130.1 The Subsystem Organization of the Sense of Smell ................... 63130.2 The Vomeronasal System................................................ 63330.3 The Septal Organ ........................................................ 63830.4 The Grüneberg Ganglion................................................ 63930.5 Noncanonical Olfactory Signaling Pathways

in the Main Olfactory Epithelium ...................................... 63930.6 Olfactory Subsystems in Humans? ..................................... 64130.7 Glossary .................................................................. 642References ....................................................................... 644

31 Disrupted Odor PerceptionThomas Hummel, Basile N. Landis, Philippe Rombaux ...................... 65331.1 Epidemiology of Olfactory Loss ......................................... 65331.2 Definitions of Olfactory Dysfunction ................................... 65431.3 Otorhinolaryngological Examination .................................. 65431.4 Questionnaires........................................................... 65731.5 Psychophysical Methods of Olfactory Testing.......................... 65731.6 Electrophysiological/Imaging Techniques ............................. 65831.7 Causes and Symptoms of Smell Disorders ............................. 65931.8 Symptoms/Quality of Life ............................................... 66131.9 Spontaneous Recovery of Smell Disorders ............................. 66131.10 Treatment of Olfactory Disorders ....................................... 66231.11 Concluding Remarks and Outlook...................................... 663References ....................................................................... 664

32 Human and Animal Olfactory Capabilities ComparedMatthias Laska .................................................................. 67532.1 Olfactory Sensitivity ..................................................... 67832.2 Olfactory Discrimination Ability ........................................ 68132.3 Qualitative Comparisons of Olfactory Capabilities Between Species. 684

Contents XXVII

32.4 General Conclusions..................................................... 685References ....................................................................... 686

33 Ectopic Expression of Mammalian Olfactory ReceptorsSophie Veitinger, Hanns Hatt ................................................... 69133.1 Discovery of Ectopically Expressed Olfactory Receptors............... 69133.2 Functionality of Ectopically Expressed ORs ............................ 69233.3 Olfactory Receptors as Cell-Cell Recognition Molecules .............. 69333.4 Olfactory Receptors in Migration and Pathfinding Processes ........ 69333.5 Diagnostical and Therapeutical Potential of Olfactory Receptors .... 69333.6 Endogenous Ligands for Ectopically Expressed Olfactory Receptors . 70133.7 Signaling Pathways of Olfactory Receptors in Non-Olfactory Tissues 70133.8 Is it Really Ectopic Expression? ......................................... 70333.9 Challenges and Future Prospects of Ectopic OR Research ............ 703References ....................................................................... 704

34 Spices and Odorants as TRP Channel ActivatorsKristina Friedland, Christian Harteneck ....................................... 71134.1 TRP Channels – Olfaction, Chemesthesis and Taste................... 71134.2 Secondary Plant Compounds Activating TRP Channels ............... 71434.3 Health Benefits of Spices ............................................... 71534.4 Negative Effects of TRP Channel Activators ............................ 71734.5 Conclusion ............................................................... 717References ....................................................................... 718

35 Anti-Inflammatory Effects of Odor CompoundsJessica Walker, Veronika Somoza ............................................... 72335.1 Relevance of Identifying Anti-Inflammatory

Active Odor Compounds................................................. 72335.2 Mechanisms Underlying the Anti-Inflammatory Activity ............ 72535.3 Assessing the Anti-Inflammatory Activity of Odor Compounds...... 72735.4 Identification of Anti-Inflammatory Odorants........................ 72835.5 Risk Assessment ......................................................... 72935.6 Summary and Outlook .................................................. 730References ....................................................................... 730

36 Skin Sensitization of Odorant MaterialsAndreas Natsch, Graham Ellis .................................................. 73536.1 Molecular Mechanism Behind Skin Sensitization ..................... 73636.2 Structural Classes of Sensitizing Molecules

with Special Emphasis on Odorants ................................... 73736.3 Testing for Skin Sensitization ........................................... 74036.4 Epidemiology of Allergy to Odorants................................... 743References ....................................................................... 745

37 Aroma Therapy in NeonatologyMichael Thiel..................................................................... 74937.1 Data on Aroma Therapy in Neonatology ............................... 75037.2 Possible Explanations ................................................... 75237.3 Neonatology-Related Research and Aroma Therapy ................. 753References ....................................................................... 753

XXVIII Contents

Part E Psycho-Physical and Cognitive Aspectsof Odor Perception

38 Cortical Olfactory ProcessingJessica Freiherr................................................................... 75938.1 Sniffing of Odors......................................................... 75938.2 Olfactory Epithelium .................................................... 76038.3 Olfactory Bulb............................................................ 76138.4 Central Olfactory Pathways and Networks ............................. 76138.5 Conclusion ............................................................... 764References ....................................................................... 764

39 Behavioral and Neural Determinantsof Odor Valence PerceptionJanina Seubert, Christina Regenbogen, Ute Habel, Johan N. Lundström ... 76939.1 Odor Valence and Behavior............................................. 77039.2 Determinants of Odor Valence.......................................... 77339.3 Valence Coding in the Human Brain ................................... 77739.4 Conclusion ............................................................... 780References ....................................................................... 781

40 Odor and EmotionSylvain Delplanque, Géraldine Coppin, David Sander ........................ 78740.1 Emotion as a Multicomponential Phenomenon ...................... 78840.2 Basic Emotions .......................................................... 78840.3 Dimensional Approaches ............................................... 78940.4 Beyond Valence: The Case of Feelings ................................. 79040.5 From Traditional Models to Appraisals................................. 79240.6 Functions of Emotions in Olfaction .................................... 79340.7 Conclusion ............................................................... 794References ....................................................................... 794

41 Aversive Olfactory ConditioningValentina Parma, Donald Wilson, Johan N. Lundström ...................... 79941.1 The Anatomy of Neural Circuits Involved in Mammalian

Aversive Olfactory Conditioning ........................................ 80141.2 Aversive Olfactory Conditioning-Induced

Structural and Functional Plasticity.................................... 80341.3 Clinical Applicability .................................................... 80941.4 Conclusions .............................................................. 811References ....................................................................... 812

42 Odor-Based Context Dependent MemoryMaria Larsson, Artin Arshamian, Cornell Kärnekull........................... 82142.1 Odor-Based Context-Dependent Memory ............................. 82142.2 Autobiographical Odor Memory ........................................ 823References ....................................................................... 825

43 Infants and Children Making Sense of ScentsBenoist Schaal ................................................................... 82743.1 Nasal Chemosensation in the Fetus.................................... 82843.2 Newborn Infants ........................................................ 830

Contents XXIX

43.3 From Preverbal Infants to Prepubertal Children ...................... 83543.4 Conclusions and Prospects.............................................. 841References ....................................................................... 842

44 Olfaction and Eating BehaviorSanne Boesveldt ................................................................. 84944.1 Metabolic Influences on Odor Perception ............................. 85044.2 Anticipation Effects of Odor Exposure ................................. 85144.3 Consumption ............................................................ 85244.4 Effect of Retronasal Odors on Eating Behavior ........................ 85344.5 Methodological Considerations ........................................ 85544.6 Conclusion ............................................................... 856References ....................................................................... 856

45 Olfaction and SleepOfer Perl, Anat Arzi, Ilana S. Hairston, Noam Sobel .......................... 86145.1 Sleep: Architecture and Measurement................................. 86145.2 Sleep Mechanisms....................................................... 86445.3 Olfactory Stimulation, Unlike Most Forms

of Sensory Stimulation, Is not Sleep-Perturbing ..................... 86645.4 Odors May Act as Sleep-Promoters:

From Aromatherapy to Neuroscience .................................. 86745.5 Processing of Olfactory Information in Sleep.......................... 86845.6 The Olfactory System and the Wake-Sleep

Regulation System are Highly Connected.............................. 87045.7 Olfaction in the Study of Sleep Mentation ............................ 87045.8 Olfaction and Sleep Disorders .......................................... 87345.9 Final Words .............................................................. 874References ....................................................................... 874

46 The Intranasal Trigeminal SystemJohannes Frasnelli, Simona Manescu.......................................... 88146.1 General Neuroanatomy of the Trigeminal System .................... 88246.2 Chemosensory Trigeminal Neuroanatomy ............................. 88346.3 Trigeminal Perception................................................... 88346.4 Assessment of the Trigeminal System.................................. 88546.5 Interactions Between Olfaction and Trigeminal System .............. 88846.6 Clinical Aspects of the Trigeminal System ............................. 890References ....................................................................... 891

47 Cross-Modal Integration in Olfactory PerceptionHan-Seok Seo, Thomas Hummel ............................................... 89747.1 Orthonasal and Retronasal Olfaction .................................. 89847.2 Cross-Modal Correspondences Between Olfactory

and Other Sensory Cues ................................................. 90147.3 Influences of Visual Cues on Olfactory Perception .................... 90347.4 Influences of Gustatory Cues on Olfactory Perception ................ 90647.5 Influences of Auditory Cues on Olfactory Perception ................. 90847.6 Influences of Trigeminal Cues on Olfactory Perception ............... 90947.7 Influences of Tactile Cues on Olfactory Perception.................... 91047.8 Conclusion ............................................................... 911References ....................................................................... 912

XXX Contents

Part F Human Body Odor, Chemo-Communicationand Behavioral Implications

48 Analysis and Chemistry of Human OdorsChristian Starkenmann ......................................................... 92148.1 Human Sweat ............................................................ 92148.2 Volatile Organic Compounds of Human Urine ......................... 92748.3 Volatile Organic Compounds in Human Feces and Pit Latrines ...... 93048.4 Conclusions .............................................................. 933References ....................................................................... 933

49 Biochemistry and Genetics of Human Axilla OdorAndreas Natsch .................................................................. 93749.1 Importance of Bacteria in Odor Formation ............................ 93749.2 Specific Odor-Forming Bacterial Populations Colonizing the Axilla . 93849.3 Specific Odorant Precursors Secreted in the Human Axilla ........... 93849.4 Malodor Releasing Enzymes in Axilla Bacteria

and Corresponding Genes .............................................. 94049.5 Fingerprints of Released Odors in Human Individuals ............... 94249.6 Effect of the Human Leukocyte Antigen (HLA) on Precursor Release? 94349.7 Ethnic Effects on Odor Formation and the ABCC11 Polymorphism .. 94449.8 Outlook: Toward More Specific Deodorants ........................... 944References ....................................................................... 945

50 Individual Variation in Body OdorJan Havlíček, Jitka Fialová, S. Craig Roberts .................................. 94950.1 Personality ............................................................... 95050.2 Sexual Orientation....................................................... 95050.3 Hormonal Influences.................................................... 95050.4 Diet ....................................................................... 95150.5 Diseases and Disorders.................................................. 95450.6 Conclusion ............................................................... 958References ....................................................................... 959

51 Processing of Human Body OdorsValentina Parma, Amy R. Gordon, Cinzia Cecchetto,Annachiara Cavazzana, Johan N. Lundström, Mats J. Olsson ............... 96351.1 The Microsmatic Fallacy ................................................. 96451.2 Human Chemosignals ................................................... 96551.3 How do Human Senders Produce Chemosignals?..................... 96551.4 Human Axillary Chemosignals for Experimental Purposes ........... 96751.5 Central Processing of Human Chemosignals........................... 97051.6 Human Chemosignals of Harm Avoidance............................. 97251.7 Central Processing of Human Chemosignals Involved

in Harm Avoidance ...................................................... 97551.8 A Clinical Perspective on Human Chemosignals....................... 97751.9 Conclusions .............................................................. 978References ....................................................................... 979

52 Human Chemosensory CommunicationBettina M. Pause ................................................................ 98752.1 Interindividual Communication........................................ 987

Contents XXXI

52.2 Release of Chemical Substances in Humans........................... 98952.3 Perception of Social Chemosignals in Humans........................ 98952.4 Chemosensory Communication via Single Molecules ................. 99152.5 Chemosensory Communication via Complex Body Fluids ............ 99852.6 Outlook................................................................... 1002References ....................................................................... 1002

Part G Odors in Language, Culture and Design

53 Odor Descriptions from a Language PerspectiveJeannette Nuessli Guth, Maren Runte ......................................... 101353.1 Comparison of Odor Terms Used by Experts

and in Everyday Language.............................................. 101553.2 Odor Terms in Everyday Language ..................................... 101953.3 Strategies to Describe Odor Perceptions in Everyday Language...... 102353.4 Conclusion ............................................................... 1025References ....................................................................... 1026

54 The Scent Creation ProcessElise Sarrazin .................................................................... 102754.1 The Role of Perfumer, Between Craftsman and Artist ................ 102754.2 Perfumery is an Art...................................................... 103054.3 Perfumery is a Science .................................................. 103654.4 New Challenges in Scent Creation...................................... 103954.5 Conclusion ............................................................... 1040References ....................................................................... 1040

55 Odor in Immersive EnvironmentsManfred Dangelmaier, Roland Blach .......................................... 104355.1 Defining Immersive Environments ..................................... 104355.2 Virtual Reality............................................................ 104555.3 Multimodality in Immersive Environments ........................... 104655.4 Functional Aspects of Odor in Human-Computer Interaction ....... 104755.5 System Design ........................................................... 104755.6 Applications.............................................................. 1049References ....................................................................... 1050

56 Odor in MarketingNathalie Nibbe, Ulrich R. Orth .................................................. 105356.1 Using Scent for Marketing .............................................. 105356.2 Effective Characteristics and Processing of Scent ..................... 105456.3 Consumer Response to Scent ........................................... 105656.4 Cross-Modal Effects ..................................................... 105956.5 Moderators ............................................................... 106056.6 Ethical Aspects ........................................................... 106156.7 Future Research ......................................................... 106256.8 Conclusion ............................................................... 1063References ....................................................................... 1063

XXXII Contents

57 Sensual Perception in ArchitectureKatinka Temme .................................................................. 106957.1 Space and Limits of Space .............................................. 107057.2 Sensual Space ........................................................... 107157.3 Sensual Perception as a Design Method in Architecture ............. 107657.4 The Ideal Laboratory .................................................... 1077References ....................................................................... 1079

58 Microdosing of ScentsMartin Richter ................................................................... 108158.1 Microdosing of Scent .................................................... 108158.2 Generating Scent Molecules ............................................ 108558.3 Microdosing Systems: Concepts ........................................ 108758.4 New Applications for Microdosing of Scent............................ 109458.5 Conclusions .............................................................. 1097References ....................................................................... 1097

About the Authors ................................................................ 1099Detailed Contents................................................................. 1115Subject Index ...................................................................... 1137

XXXIII

List of Abbreviations

1-D one-dimensional2-D two-dimensional2-DG 2-deoxy-D-glucose3-D three-dimensional3M2H 3-methyl-2-hexenoic acid4-D four-dimensional

A

AAT alcohol acyl transferaseABC ATP-binding cassetteACC anterior cingulate cortexACD allergic contact dermatitisACP acyl carrier proteinADH alcohol dehydrogenaseADI acceptable daily intakeAEDA aroma extract dilution analysisAHI apnea-hypopnea indexALA alpha-lipoic acidALDH aldehyde dehydrogenaseAM autobiographical memoryAMP adenosine monophosphateAMP adenosine 50-monophosphateAMRE axillary malodor releasing enzymeANOVA analysis of varianceAO anterior olfactory nucleusAOB accessory olfactory bulbAOP adverse outcome pathwayAPC anterior piriform cortexAPCI atmospheric pressure chemical ionizationAPET added portions exposure techniqueAPI-MS atmospheric pressure ionization mass

spectrometryAPLI atmospheric pressure laser ionizationARP Amadori Rearrangement ProductARS acetone reference standardATP adenosine triphosphate

B

BDI Beck depression inventoryBDNF brain-derived neurotrophic factorBLA basolateral amygdalaBMI body mass indexBOLD blood-oxygen-level-dependentBOTT basic odor term testBSA bovine serum albuminBTTT basic taste term test

C

CA Cornu AmmoniscAMP cyclic adenosine monophosphateCANN cortical-based artificial neural network

CCCRC Connecticut Chemosensory ClinicalResearch Center test

CD cyclodextrinCE carboxyl esteraseCI chemical ionizationCNG cyclic nucleotide gatedCNS central nervous systemCOA conditioned odor aversionCREB cAMP responsive element bindingCSERP chemosensory event-related potentialCSIA compound-specific isotope analysisCWA chemical warfare agent

D

DAG diacylglycerolDES diethyl sulfideDESI desorption electrospray ionizationDG dentate gyrusDHE dynamic headspace extractionDMAPP dimethylallyl diphosphateDMDS dimethyl disulfideDMS dimethyl sulfideDMSO dimethyl sulfoxideDNA deoxyribonucleic acidDNPH dinitrophenylhydrazine

E

e-nose electronic noseEBC exhaled breath condensateEC entorhinal cortexee enantiomeric excessEEG electroencephalographyEESI extractive electrospray ionizationegr-1 early growth response protein 1EH epoxide hydrolaseEI electron ionizationELDI electrospray-assisted laser desorptionEMEM Eagle’s modified essential mediumEMG electromyographyEOG electroolfactogramEOS emotion and odor scaleser enantiomeric ratioEROM European reference mass odorERP event-related potentialEs-GC enantioselective gas chromatographyESP epithio-specifier proteinsEST expressed sequence tagEtSH ethanethiol

F

FAD flavin adenine dinucleotideFD flavor dilution

XXXIV List of Abbreviations

FDSS functional drug screening systemFDT flow drift tubeFFI food freshness indicatorFGE flavoring group evaluationFID flame ionization detectionFLEC field and laboratory emission cellfMRI functional magnetic resonance imagingFPD flame photometric detectionFT Fourier transformFTIR Fourier transform infraredFWHM full-width at half maximum

G

GABA � -amino butyric acidGC gas chromatographyGC-FID gas chromatography with

flame-ionization detectionGC-IRMS gas chromatography isotope ratio mass

spectrometryGC-O gas chromatography-olfactometryGC-PO GC-pedestal olfactometerGC-RO GC-recomposition-olfactometryGC-SNIF GC-surface of nasal impact frequencyGEOS Geneva emotion and odor scaleGFP green fluorescent proteinGG Grüneberg ganglionGPCR G protein coupled receptorGPMT guinea pig maximization test

H

HCC-HS high concentration capacity headspaceHD homeodomainHEK human embryonic kidney cellHLA human leukocyte antigenHLB hydrophilic-lipophilic balanceHPLC high performance liquid chromatographyHRIPT human repeat insult patch testHS headspaceHS-LPME headspace liquid-phase microextractionHS-SPME headspace solid-phase microextractionHS-STE headspace sorptive tape extractionHSSE headspace sorptive extractionHTS high-throughput screeningHVP hydolysed vegetable protein

I

IAMS ion attachment mass spectrometryIAT implicit association testIBMX isobutyl methyl xanthineIC50 half-minimal inhibitory concentrationIGF insulin-like growth factorIL interleukinIMR ion-molecule reactionIMS ion mobility spectrometryIPP isopentenyl diphosphateIR infrared

IRMS isotope ratio mass spectrometryIT ion trapIT-MS ion trap mass spectrometryITE individual threshold estimate

J

JND just noticeable difference

K

KC K-complexKegg Kyoto Encyclopedia of Genes and

GenomesKSOM Kohonen self-organizing map

L

LAB lactic acid bacteriaLAESI laser-assisted electrospray ionizationLBP LPS-binding proteinLC liquid chromatographyLDL low density lipoproteinLLE liquid-liquid extractionLLNA local lymph node assayLOD limit of detectionLOQ limit of quantitationLOT lateral olfactory tractLOX lipoxygenaseLPP late positive event-related potentialLPS lipopolysaccharide

M

MALDI-ESI matrix-assisted laser desorptionelectrospray ionization

MAP modified atmosphere packagingMCC multicapillary columnMCS multiple chemical sensitivityMDGC multidimensional GCMDR multidrug resistenceMDT mediodorsal thalamic nucleusMEG magneto-encephalogramMeSH methanethiolMEST mouse ear swelling testMHC major histocompatibility complexMICA metabolism of ingestion correlated

amountMLF malolactic fermentationMOB main olfactory bulbMoE margin of exposureMOE main olfactory epitheliummOR-EG mouse olfactory receptor proteinMOSFET metal oxide semiconductor field effect

transistorsMRI magnetic resonance imagingmRNA messenger ribonucleic acidMS mass spectrometryMSDI maximized survey-derived daily intake

List of Abbreviations XXXV

MSG monosodium glutamatemTAMDI modified theoretical added maximum

daily intakeMUFA monounsaturated fatty acidMUP major urinary protein

N

NIF nasal impact factorNMR nuclear magnetic resonanceNOAEL no observed adverse effect levelNOEL no observed effect level

O

O olfactometryOAV odor activity valueOB olfactory bulbOBP odor binding proteinOCAM olfactory cell adhesion moleculeODR odorant responseODT odor detection thresholdOE olfactory epitheliumOERP olfactory event-related potentialOET open epicutaneous testOFC orbitofrontal cortexOIRS odor intensity referencing scaleOMP olfactory marker proteinOR olfactory receptorORN olfactory receptor neuronORP olfactory receptor proteinORT odor recognition thresholdOSN olfactory sensory neuronOTH odor detection thresholdOU odor unit

P

PA proton affinityPAD pleasure, arousal, and dominancePAH polycyclic aromatic hydrocarbonsPAMP pathogen-associated molecular patternPBMC peripheral blood mononuclear cellPC piriform cortexPCA principal components analysisPCC posterior cingulate cortexPCR polymerase chain reactionPDMS polydimethylsiloxanePDO protected designation of originPE polyethylenePEA phenylethylalcoholPEEK polyether ether ketonePET positron emission tomographyPFC prefrontal cortexPFPD pulsed flame photometric detectorPFS passive flux samplerPI perceived intensityPIB polyisobutylenePID photoionization detector

PLC phospholipase CPMMA polymethylmethacrylatePP polypropylenePPC posterior piriform cortexpremox prematurely agedPS polystyrenePTFE polytetrafluoroethylenePTR proton transfer reactionPU polyurethanePUFA polyunsaturated fatty acidPUT provocative use test

Q

QC quality controlQOD questionnaire olfactory disorderQoL quality of lifeQRA quantitative risk assessment

R

RAS retronasal aroma simulatorREM rapid eye movementRIA retention index allowanceRNA ribonucleic acidROAT repeated open application testROS reactive oxygen speciesRTL retention time locking

S

S-HS static headspaceSAFE solvent assisted flavor evaporationSAR structure–activity relationshipSCFA short-chain fatty acidSDE simultaneous distillation extractionSEC size exclusion chromatographySESI secondary electrospray ionizationSID selected ion detectionSIDA stable isotope dilution analysisSIFDT selected ion flow drift tubeSIFT selected ion flow tubeSIM selected ion monitoringSIRA stable isotope ratio analysisSND sinunasal diseaseSNP single nucleotide polymorphismSOA secondary organic aerosolSPE solid phase extractionSPME solid phase micro extractionSrc sarcomaSVHC substance of very high concernSVOC semivolatile organic compound

T

TA titratable acidityTAAR trace amine associated receptorTAGA trace atmospheric gas analyzerTD thermal desorption

XXXVI List of Abbreviations

TDS temporal dominance of sensationtERP trigeminal event-related potentialTIC toxic industrial compoundTIR Toll/interleukin-1 receptorTLR Toll-like receptorTM transmembraneTMR targeted memory activationTNF tumor necrosis factorTOF time-of-flightTPOA taste potentiated odor aversionTPS terpene synthaseTRP transient receptor potentialTRPC transient receptor potential channelTSS total soluble solidsTu olfactory tubercleTVOC total amount of volatile organic

compounds

U

UPSIT University of Pennsylvania smellidentification test

URTI upper respiratory tract infectionUTA untypical aging off-flavorUV ultravioletUV–Vis ultraviolet-visible

V

V-SMOW Vienna Standard Mean Ocean WaterVMR volume mixing ratioVNO vomeronasal organVOC volatile organic compoundvPvB very persistent and very bioaccumulativeVR vomeronasal receptorVSC volatile sulfur compoundVSN vomeronasal sensory neuronVUV-SPI vacuum ultraviolet single-photon

ionization

W

WOF warmed-over-flavor