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“blue”glomeruli).Efferent(bulbofugal,blue)fibersprojectfrommitral/tuftedcellstosecondaryolfactorystructures;afferent(centrofugal,
gray)fibersprojecteitherfromcontralateralmitralcellsoripsilateralcentralnucleiandsynapsetoglomerularorgranulecells,respectively.
b,cImmunohistochemicalfeaturesofthemouse(b)andhuman(c)olfactorybulb(OB).Arabicnumbersagainindicatethecorticalstructure
(see(a)).DoubleimmunofluorescenceofthemouseOB(b)usingantibodiesagainstOMP(green)andthepan-neuronalmarkerPGP9.5
(red)showsamoredistinctlaminationthaninan80-year-oldhuman.OMP,reactiveneuronsorglomeruli;G,glomerulus;M,mitralcell.
CellLayers
Asmentionedabove,theOBshowsacorticallamination.Itmainlyconsistsoftheolfactorynervelayer,
theglomerularlayer,theinternalplexiformlayer,themitral/tuftedcelllayer,theexternalplexiformlayer,
and the granule cell layer (Fig. 3.4). ORNs assemble on the OB´s surface and immerse in easily
detectable spherical structures, glomeruli, in which they connect with dendrites of mitral cells.
Glomeruliaresurroundedbytwomajorgroupsofinhibitoryinterneurons,periglomerularcellsandshort
axon(SA)cells,whichareresponsibleforfinetuningolfactoryinformationmediatedthroughmitralcells.
The deepest cortex layer consists of columnar arrangements of another cell type—granular cells—
derived from precursor neurons that migrate from the rostral migratory stream and the subventricular
zone. The regenerative capacity of those interneurons is extraordinarily high. Genetic tracer studies
suggestthearrivalof~30,000newneuronsperdayintheOB.
10
However,onlyasubsetoftheseneurons
climbsuptoconverttotheinterneuronsmentionedabove.
11
BothkindsofOBinterneuronsarealsotarget
neuronsforinhibitoryprojectionsfrom the contralateral anterior olfactory nucleusthroughthe anterior
portionoftheanteriorcommissure.
Convergence
Fromrodentstudiesitisknownthatseveralolfactoryneuronsexpressingthesamereceptorconvergeonto
two glomeruli. In consequence, a given odorant elicits a characteristic spatial pattern of glomerular
activities,
12,13
initiatinganodor-specificactivationpatternalreadyattheOBlevel.Odor-relatedspatial
positions are conserved across different individuals of the same species. However, this topographic
organizationhasnotyetbeendemonstratedinneocorticalbrainareas.
Processes of GABAergic periglomerular cells project to a single glomerulus for intraglomerular
processing, whereas GABA- and dopaminergic SA cells extend to multiple glomeruli, thus providing
interglomerularcrosstalk.
14
Localcircuitsamonglargeprojectionneurons(mitral/tuftedcells)andsmall
OBinterneurons mediate lateral inhibitionamong glomerular units to enhance contrastrelatedto odor
informationthatisforwardedtohigherorderolfactorystructures.
2
The olfactory bulb is the first relay for incoming olfactory information. Lateral inhibition and
convergenceattheleveloftheolfactorybulbseemtobeimportantforcontrastingdifferentolfactory
perceptionsagainsteachother.
WhatisSpecialabouttheHumanOlfactoryBulb?
Comparedwithlaboratoryanimals,thevolumeofthehumanOBismuchsmallerrelativetototalbrain
size (Figs. 3.3 and 3.5). Furthermore, data obtained from 42 human autopsies (mean age 77 years)
revealeda remarkable interindividualvariability inOBsize(mean:83µL;range:50 to142µL).
8
As
mentionedabove,magneticresonanceimaging(MRI)studiesrevealedastrongcorrelationbetweenOB
sizeandolfactoryperformance.
6
However,preciseinformationonwiring,presortingofindividualORNs
in the periphery, and receptor-specific glomerular unit organization has not yet been determined in
humans.
Besides this, the number of glomeruli in thehuman OB is considerably higher than animal models
wouldpredict.AnimalstudiesprovideevidencethatORNsexpressingoneparticularreceptorprojectto
onlytworeceptor-specificglomeruli.This2:1model(twoglomeruliperreceptor)assumesthatif1,000
functionalreceptorgenesexist(asinmice),thereare2,000glomeruliperOB.
15
Inhumans,however,who
haveonly~340to400functionalreceptorgenes,
16,17
onemight predict680 to800 glomeruli, whichis
clearlycontradictedbyhistomorphologicalestimationsrangingfrom2,975to9,325glomeruliperhuman
OB.
18,19
Thus the 2:1 ratio originating from other mammals is greater by a factor of eight (16:1) in
humans. These results suggest that the convergence described for rodents or other mammals mightbe
distinctfromthatinhumans,pointingtoafundamentaldifferenceinhumanOBorganization.
19
ProjectionTargetsofMitralCellsintheOlfactoryBulb(“SecondaryOlfactoryStructures”)
•Anteriorolfactorynucleus
•Anteriorperforatedsubstanceandunderlyingventralstriatum(nucleusaccumbens)
•Piriformcortex
•Periamygdalarcortex/medialamygdala
•Entorhinalcortex
Fig.3.5a,bOlfactorybrainregionsintheorbitofrontalareainaschematicdrawing(a)andoverlaidontoastructural,T2-weightedmagnetic
resonanceimage(b). Thebluelinedelineates theolfactory epitheliumin theupperturbinateofthehumannose(aOF, anterior;lOF,lateral;
mOF,medialorbitofrontalgyrus;GR,rectalgyrus).(Fig.3.5aadaptedfromSchünkeetal.2007.
66
)
AnteriorOlfactoryNucleusasInhibitorofContralateralInformation
TheolfactorypedunclecontainsmitralcellaxonsbundledintheLOTandissituatedlaterallytotherectal
gyrusandadjacenttotheolfactorysulcusofthebasalforebrain(Fig.3.2).Thetermolfactorypeduncleis
usuallyusedfordescribingtheAONandtheanteriorpartoftheLOTasawhole.
TheAONisapredominantlytwo-layeredstructurecomposedofneuronsscatteredacrossthecourseof
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the olfactory peduncle. It is the major relayforfibers to andfrom thecontralateral OB.In contrastto
ipsilateralconnections,contralateralneuropeptideYprojectionsthroughtheanteriorcommissureseemto
decreaserapidlyafterbirthincats.
20
ThesignificanceoftheAONforprocessingofolfactorystimuliis
unclear.ItisknownthattheAONmainlycontainsinhibitoryfiberstoenhancethecontrastofipsilateral
olfactory information. Notably, the AON is oneof the first nuclei affected in patients withParkinson
diseaseormultiplesystematrophy.
21–23
AnteriorPerforatedSubstanceandUnderlyingVentralStriatumMediatethe
InductionofSexualArousalviaOlfactoryCues
Theolfactorytubercle,aprominentstructureinanimals,israrelydistinguishableinhumans.Instead,the
anteriorperforatedsubstance(APS)mightbeconsideredasitshumanhomologue(Fig.3.2a).TheAPSis
athree-layeredcortexwithcharacteristiccellclusters,theislandsofCalleja.ThefunctionoftheAPSin
humans is largelyunknown. The ventral parts of the caudate nucleus, putamen, pallidum,and nucleus
accumbens(brainareas that are situated adjacent tothe APS) are conjointlyreferred toasthe ventral
striatum (Fig. 3.2b). As recently reviewed by Heimer's group,
3,24
animal studies point to a direct
projectionpathwayfromtheolfactorycortextowardthebasalganglia(ventralstriatum)ratherthantothe
hypothalamus.The functionalsignificanceof this projectionprobablyinvolves themediation ofsexual
arousalelicitedbylearnedolfactorycues.
5
DifferentLevelsofCorticalProcessingTakePlaceinthePiriformCortex
Thepiriformcortex(pirC) isathree-layered,pearshapedallocorticalstructurespanningthejunctionof
themedialfrontalandtemporallobes.Itisthemainandmostimportantprojectionareaofmitral/tufted
cell axons inthe OB (Fig. 3.2). From here, projections terminatein subsequent subcortical areas via
allocorticostriatal pathways into the ventral striatum and hypothalamus. Further neocortical olfactory
areasincludetheinsulaandorbitofrontalgyri(seebelow).
Earlyolfactory neuroimaging experimentsrevealedthatthepirC is involvedinbasicprocessing of
passiveolfactorystimulation.
25–28
However,therecentdevelopmentofmodernolfactometry,aswellas
functionalneuroimagingmethodsandstudydesigns,hasenabledresearcherstoprovide further insights,
leadingtoachangeofconceptregardingthefunctionofthepirC.IthasbeenshownthatthepirCcanbe
subdividedintoananterior,orfrontal,andaposterior,ortemporal,portion,and,althoughhistologically
similar, these portions show a functional heterogeneity.
29
They are involved in different olfactory
processingsteps.Whiletheanteriorportionisconsideredprimarilytobeinvolvedinbasicorlowlevel
olfactoryprocessingsuchaspassivesmellingandencodingofthechemicalormolecularstructureofan
odorant,theposteriorportionisratherresponsibleforhigherorderolfactoryprocessingsuchashedonic
quality evaluation or discrimination and encoding of the perceptual character of an odorant.
30–33
The
pirC,asaprominenttargetstructureofOBafferents,isalsocriticallyinvolvedinolfactorymemoryand
learningaswellasmultisensoryintegrationprocesses.Theresultsofarecentmeta-analysis
34
implythat
thepiriformcortexisconsistentlyactivatedacrossasetofolfactoryfunctionalimagingexperiments(Fig.
3.6).
TheAmygdalaEncodesBehavioralSalienceofanOdor
Olfactory projections pass either directly or via pirC to a superficially located subdivision of the
amygdala,theperiamygdalar complex, andtothe anteriorcorticalnucleusofthe amygdala (Fig. 3.2a).
Deepersubnuclei arenot directlylinked toLOTprojections.In animals,amygdalar structuresnotonly
coordinatethemediationofaggressiveandpredatorbehavior,butalsoarelinkedtosexualdevelopment
andolfactoryrecognitionofoffspring.
35
Amygdalarnucleiarealsocanonicaltargetsofvomeronasalinput
in most animal species.
36
As the existence of a vomeronasal system in humans is questionable (see
Chapter 4), stimuli processed via the olfactory system may contribute to gender-specific reactions to
odorants or pheromone-like substances.
37
It is known that the anterior cortical nucleus and the
periamygdaloidcomplexarehighlyresponsivetoodorants.
36
Someauthorshaveconnectedthefunctionof
thecentromedialamygdalarnucleiwiththatoftheBNST,
38
ahypothalamicnucleusinvolvedinaneuronal
circuitalsoinvolvingtheOB,anteriorpiriformcortex,andpreopticarea.
Fig.3.6 Theresultsof ameta-analysisoffunctional imagingstudies usingthetechniqueofactivationlikelihood estimation.
34
Depicted are
brain areas thatare commonly recruitedowing totrigeminal stimulation(red) andpureolfactory stimulation (blue). Typicalolfactorybrain
areasarethepiriformcortex(bilaterallyin slice−14),orbitofrontalcortex(bilaterallyinslice−9), andinsula(bilaterallyinslice 6).(Adapted
fromAlbrecht2010.
34
)
Severalresearchteamsusedfunctionalimagingtechniquestoanswertheimportantquestionofwhether
theamygdalaisinvolvedincodingodorintensityorvalence.
39–42
Theresultsoftheirstudiessuggestthat
theamygdalaencodesthebehavioralsalienceofanolfactorystimulus,andthusacombinationofintensity
andvalenceisimportant.
43
Thereisalsoevidencethattheamygdalaisinvolvedinassociativelearning
alongwitholfactorymemory.
29
EntorhinalCortex:AGateforOlfactoryMemory
TheentorhinalcortexisthemostdistanttargetareaofLOTaxons(Fig.3.2)andconstitutesatransition
zone from three-layered allocortex into sixlayered neocortex. Its strongest projection turns into the
hippocampal formation. Therefore, it is assumed that the entorhinal cortex is involved in olfactory
memory processes. Results of a functional MRI (fMRI) study reveal that activation of the entorhinal
cortexcorrelateswithperceivedintensityofolfactorystimuliandthusisconsideredtobeinvolvedina
gatingprocess of intense olfactory stimuli tothe hippocampus.
44
Theentorhinal cortex shows specific
neuropathologiesinAlzheimerdisease.
45
ProjectionTargetsofSecondaryOlfactoryStructures
•Insula
•Orbitofrontalcortex
•Ventralstriatum(e.g.,nucleusaccumbens)
•Hippocampus
•Septalnuclei
•Hypothalamus(bednucleusofstriaterminalis,preopticarea)
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TheInsulaisResponsibleforIntegrationofDifferentChemosensoryStimuli
Theinsulaisanagranularcorticalareaburiedinthedepthsofthelateralsulcusandoverlaidbyopercula
(“lids”) from the frontal, parietal, and temporal lobes (Fig. 3.2). The anterior part of the insula in
particularisconsideredtobetheprimarytastecortex.Nevertheless,inrats,theinsulareceivesolfactory
projectionseitherdirectly fromthe pirC orindirectlythrougha thalamic relay.
46
In humans,fMRIand
positronemissiontomography(PET)studiesshowactivationoftheanteriorinsulainrelationtoodorant
stimulation.
47
Thepreciserole oftheinsulainodorprocessingremainssomewhat unclear.Resultsofa
recentmeta-analysis support the view thatthe insula playsanimportant role during the integration of
olfactionwithotherchemosensorymodalities.
33
Theauthorsreviewedscientificliteratureandsubmitted
the results of all functional neuroimaging experiments utilizing (1) olfactory, (2) trigeminal, and (3)
gustatorystimulationto an activation likelihood estimation(ALE) meta-analysis. A conjunctionof the
threeALEmapsrepresentingthebrainareasthatarerecruitedacrossallstudiesinthethreemodalities
revealsthatapartoftheanteriorinsulaisrecruitedduringallthreeconditions(Fig.3.7).Thusitseems
highlylikelythattheanteriorportionoftheinsulaisresponsibleforintegrationofdifferentchemosensory
experiences.
TheOrbitofrontalCortexistheIntegrationSitewithOtherSensoryModalities
Theorbitofrontalgyriinterposedbetweentherectalgyrusmediallyandtheinsulalaterallyareconsidered
tocomposetheorbitofrontal cortex(OFC;Figs.3.2a and3.5).Itconsistsofanagranular (fivelayered)
cortexandreceivesdirectinputmainlyfromthepirCandindirectinputviathethalamus.Althoughdata
fromtracerstudiesinhumansarenotavailable,reportsfrommonkeystudiessuggestaveryearlystageof
olfactory informationprocessing, inconjunction withadjacent gustatoryandvisualprocessing areas in
frontalportionsoftheOFC.
29,48
Projectionsfromandtootherbrainregionsareextensive,especiallyfor
integrationofseveral sensory modalities. Theintegrative functionofthe OFCis probablyinvolved in
generating the sensation of flavor associated with different foods.
48,49
The OFC is considered to be
involved in olfactory working memory and discrimination processes.
50
In addition, recognition and
rewardvalueofanodorisrepresentedintheOFC.
51,52
Fig.3.7Resultsofarecentactivationlikelihoodestimation(ALE)meta-analysis
33
investigatingcommonbrainactivationacrossallolfactory,
trigeminal, andgustatoryfunctional imagingstudies. Theconjunctionofall threeALE mapsrevealsthattheanteriorportionoftheinsulais
importantfor processing of stimuli inall three modalities (yellow). Thus it is suggested that the anterior insulais involvedin multisensory
integrationprocesses.(AdaptedfromLundströmetal.2011.
33
)
Forillustrationofcorticalprocessesduringsmellperceptioninhumans,imagingtechniquessuchas
fMRI,electroencephalography, or PET, in combinationwithconstant flow olfactometry, are used.
The development of sophisticated statistical analysis tools such as voxel-based morphometry,
independentcomponentanalysis,dynamicalcausalmodeling,andALEwarrantsfurtherinsights.
Non-SpecificOlfactoryBrainRegions
Functionalimagingrecordsrevealthatmanyotherbrainareasareactivatedbyolfactorystimuli,withouta
direct anatomical link to primary olfactory structures.
53
The pathways appear diffuse and include
neocorticalareasaswellas,forexample,thecerebellum,whichseemstoplayaroleinmotorcontrolof
olfaction.Onehypothesissuggeststhatsniffingactivityisregulatedbyafeedbackmechanismusingdata
onodorconcentration.
53
FunctionalCharacteristicsofOlfactoryProcessing
TheRoleoftheThalamusinOlfaction—MoreImportantthanPreviouslyThought
Common anatomical work in animals and humans implies no direct thalamic activity during central
olfactoryprocessing.CanonicalpathwaysincludeonlytworelaysfromORNsviamitral/tuftedcellsinto
thepiriformcortex,andfromtheredirectlyintomainlymedialandlateralOFCprojectionareaswithout
furthersynapse.
However,severalanimalmodelssuggestatleastoneindirectconnectionbetweenthepiriformcortex
andneocorticalareas(OFC)viathemedioposteriorthalamicnucleus,butthisrouteseemslessimportant
thandirectpathways,asitcontainsconsiderablyfewerfibers.
54
Thefirstevidenceofamoreimportant
thalamic interaction with odor processing was found during an fMRI study
55
and recently during an
investigationutilizingmultisensorystimulation.
4
Bothstudiessuggestthatattentiontosmellsenhancesthe
participation of the thalamocortical pathway, which appears to be a modulatory target of olfactory
attentionalprocessing.
Astothenecessityofathalamicinterneuroninolfaction,onemightassigncomputationalissuessuch
asconvergenceandsortingoutolfactoryinformation(“bottleneckfunctions”)askeytasksoftheOBand
thuscompareitsactivitytothatofthethalamusinothersensorysystems.
56
Task-specificOdorProcessing
The number and size of secondary olfactorystructures is tremendous,including theolfactorynucleus,
corticalnucleiofamygdala,periamygdalarcomplex,anteriorandposteriorpirC,entorhinalarea,andthe
anteriorperforatedsubstancewithadjacentventralstriatum(ortheolfactorytubercleasitshomologuein
animals),nottomentionalltertiaryneocortical,diencephalic,andbrainstemareastargetedbyprojections
of the former regions. One should wonder if, after sniffing a simple odor, the entire brain might be
involved and occupied with processing this particular sensation. Certainly, it is not. An approach to
determine thespecialization of diverse olfactory-related brain areas was introduced by measuring the
effects of task-specific stimulation. There are, for example, hemispheric differences in perception of
familiar/unfamiliar or pleasant/unpleasantstimuli. The lefthemisphere is more engaged in processing
familiar odors, and the right hemisphere in that of unpleasant odors.
57
Further, olfactory detection,
discrimination, andmemoryrecoveryfrom experience implies the hierarchical recruitmentofdifferent
structuralandfunctionalrelayswithintheolfactorysystem.
58
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Lateralization
Aspreviouslyintroduced,thereissomeevidenceforolfactorylateralization(i.e.,specificolfactorytasks
arerepresentedinonehemisphere).olfactorynervefibersprojectipsilaterallyuptothelevelofthepirC
(seeabove).Theanatomicalsubstrateforlateralizedcorticalodorantprocessingseemstobetheanterior
partoftheanteriorcommissure,butproofofthispossibilityisstilllacking.Arecentlypublished,event-
relatedsourceimagingstudyprovidesevidencethatolfactoryinformationisprocessedindifferentsteps,
startingwithipsilateralactivationoftheamygdala,parahippocampalgyrus,superiortemporalgyrus,and
insula.Subsequently,thecorrespondingcontralateralstructuresareactivatedand,finally,frontalareasof
bothhemispheresareinvolved.
59
Accordingly,monorhinallypresentedodorantsactivatebilateralhigher
orderbrainareasindependentofthesideofpresentation.
57,59
InpatientswithParkinsondisease,neuronal
activity in the right amygdala and hippocampus seems to be diminished, and it is assumed that the
selectiveimpairmentwithinthesebrainareascausestheolfactorydysfunction.
60
Inouropinion,olfactory
lateralizationeffectsshouldbecarefullyconsidered in the contextofthe interconnectivityofthenasal,
oral,andpharyngealcavities,whichrendersapureunilateralolfactorystimulationalmostimpossible.
Adaptation–Habituation
Adaptation
Intheperiphery,along-lastingolfactorystimulusleads toadecreased responseofORNs.Intracellular
calciumions(Ca
2+
)interactwithcalmodulinandcalmodulin kinaseII. AssoonastheCa
2+
/calmodulin
complex has bound to cyclic nucleotide-gated channels, the latter decrease their sensitivity to cyclic
adenosine monophosphate (cAMP) and close again. This results in a diminished number of action
potentialsdespiteasuccessfulligand–receptorinteraction,whichisequaltoadaptation.Thecontribution
oftheenzymephosphodiesterase-1c2,whichcatalyzescAMP,hasbeenrecentlyquestioned.
61
Short-term
adaptationisdefinedasadecreaseinexcitabilityduringashortodorpresentationwithinafewseconds.
Also,alongeradaptation(overminutes)hasbeenobserved.
62
Thedurationofadaptationdependsonthe
activityoftheNa
+
/Ca
2+
exchangercomplex,whichtransportsCa
2+
outofthecell.Adaptationoccursin
peripheral ORNs, but it has been shownthat desensitization is smaller than the decrease of intensity
estimates.
63
The pirC might therefore play a more important role in adaptation than the OB or the
olfactoryreceptors.ThepirCprocessesinformation frommitralcellsoftheOBandmightreactwitha
temporarysynapticdepression(corticaladaptation).
64
Habituation
In contrastto peripheral or central adaptation,the term habituation implies a more complex, central,
multineuronallearningprocess, whichprotectsthe organism against persistent,“non-important” signals
fromitsenvironment.Itisdependentontheinputofolfactorysignals,butalsoemploysaseriesofnon-
olfactory circuits, such as brainstem nuclei involved in cardiovascular regulation.
65
This phenomenon
(“sensorygating”)maybetriggeredbyvariousstimuliorbycorticaldesensitization.
The piriform cortex seems to play a more important role in adaptation processes than olfactory
receptorcellsortheolfactorybulb.
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4TheHumanVomeronasalSystem
MartinWitt,MichaelMeredith
Oneofthemostintriguingchemosensorysubsystemsinhumansisthe“vomeronasalorgan”(VNO).Since
its discoveryin man by Frederic Ruysch in1703
1
andmore systematic observations indomesticated
animals by Ludwig Lewin Jacobson in 1811, few organs have undergone more illustrious functional
interpretationsthan the VNO.One of the most common misunderstandings concerning its functionality
reflects inappropriate analogies with nonhuman vertebrates with respect to influences on social and
sexualbehavior.
In many nonhuman mammals, the VNO is a complex ofdifferent component structures that detects
specific chemical communication signals (commonly referred to as pheromones) and conveys the
informationtothecentralnervoussystem.
TheVomeronasalOrganisFunctionalinManyNonhuman
VertebratesandhasDefinedComponents
The VNO is present in most amphibians, reptiles, and mammals, but is lacking in some successful
phylogenetic groups, such as crocodiles, birds, or marine mammals. The anatomy of the VNO varies
considerablyamongthedifferentclasses.Exceptinamphibians,theVNOisanatomicallyseparatedfrom
the main olfactory epithelium.Itis composed ofdifferenttissuesandentities, such asthe vomeronasal
duct(VND),seromucousglands,thevomeronasalnerve,avomeronasalcapsule,andavenouspumping
system (Fig. 4.1).
2
The VND in rodents is a blind-ending channel running in the lower nasal septum
parallel to the floor ofthe nasal cavity.On its medial side it is lined with a pseudostratified sensory
epitheliumthatcontainssensoryneurons,supportingcells,andbasalcells,acellularorganizationsimilar
tothatofthemainolfactoryepithelium.Thevomeronasalepithelium(VNE)ismuchthickerandcontains
two maintypesofvomeronasal sensory neurons,whichexpress two familiesofvomeronasal receptor
genes: guanosinetriphosphate–bindingproteins(G-proteins);andTRPC2transduction-channelproteins.
These, and olfactory marker protein (OMP, Fig. 4.1b), are components of the accepted (canonical)
vomeronasal transduction pathway, buttheyare missing inhumans. The axons ofvomeronasal sensory
neuronsprojecttotheaccessory olfactory bulb(AOB),
3
whichisalso missing in humans.AOBoutput
passes principally tothemedialnucleusoftheamygdala,whichintegrateschemosensoryandhormonal
information,
4
and is thought to interpret the meaning of chemical signals.
5
Outputs from the medial
amygdala, and from other nuclei receiving independent AOB input, are directed to hypothalamic and
preopticnuclei,thecontrolcentersforneuroendocrinefunctionsandsocialbehavior.
6–8
Severalvolatile
andnonvolatile compoundsinmouseurineandvariousskin secretionsstimulatespecificVNOsensory
neurons,
9
butdatalinkingparticularligandstospecificreceptorsarestillscarce.
AnatomyoftheVomeronasalSysteminHumans
PrenatalDevelopmentandRegressioninHumans:“Classical”VomeronasalOrgan
StructureSuggestsPossibilityofPrenatalFunction
Development
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