Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Region
HypothalamusWikipedia,thefreeencyclopedia
Area
Nucleus
Function[7]
Preoptic Preopticnucleus
Anterior
Medialpreopticnucleus
Regulatesthereleaseofgonadotropichormonesfromtheadenohypophysis
Containsthesexuallydimorphicnucleus,whichreleasesGnRH,differentialdevelopment
betweensexesisbaseduponinuterotestosteronelevels
Supraopticnucleus
Vasopressinrelease
Oxytocinrelease
Medial Paraventricularnucleus
thyrotropinreleasinghormonerelease
corticotropinreleasinghormonerelease
oxytocinrelease
vasopressinrelease
somatostatinrelease
Anteriorhypothalamicnucleus
thermoregulation
panting
sweating
thyrotropininhibition
Suprachiasmaticnucleus
Circadianrhythms
Lateral Lateralnucleus
Dorsomedialhypothalamic
nucleus
https://en.wikipedia.org/wiki/Hypothalamus
SeeLateralhypothalamus#Functionprimarysourceoforexinneuronsthroughoutthebrainand
spinalcord
BloodPressure
HeartRate
GIstimulation
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Medial Ventromedialnucleus
satiety
neuroendocrinecontrol
Tuberal
Arcuatenucleus
Lateral
Lateralnucleus
Growthhormonereleasinghormone(GHRH)
feeding
Dopaminemediatedprolactininhibition
SeeLateralhypothalamus#Functionprimarysourceoforexinneuronsthroughoutthebrainand
spinalcord
Lateraltuberalnuclei
Mammillarynuclei(partof
mammillarybodies)
memory
Posteriornucleus
Increasebloodpressure
pupillarydilation
shivering
vasopressinrelease
Medial
Posterior
Lateralnucleus
Lateral
Tuberomammillarynucleus[8]
SeeLateralhypothalamus#Functionprimarysourceoforexinneuronsthroughoutthebrainand
spinalcord
arousal(wakefulnessandattention)
feedingandenergybalance
learning
memory
sleep
Seealso:ventrolateralpreopticnucleus,periventricularnucleus.
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Acrosssectionofthe
monkeyhypothalamus
displays2ofthemajor
hypothalamicnucleion
eithersideofthefluid
filled3rdventricle.
Hypothalamicnuclei
Hypothalamicnucleion
onesideofthe
hypothalamus,shownin
a3Dcomputer
reconstruction[9]
Neuralconnections
Thehypothalamusishighlyinterconnectedwithotherpartsofthecentralnervoussystem,inparticularthebrainstemanditsreticularformation.Aspart
ofthelimbicsystem,ithasconnectionstootherlimbicstructuresincludingtheamygdalaandseptum,andisalsoconnectedwithareasofthe
autonomousnervoussystem.
Thehypothalamusreceivesmanyinputsfromthebrainstem,themostnotablefromthenucleusofthesolitarytract,thelocuscoeruleus,andthe
ventrolateralmedulla.
Mostnervefibreswithinthehypothalamusrunintwoways(bidirectional).
Projectionstoareascaudaltothehypothalamusgothroughthemedialforebrainbundle,themammillotegmentaltractandthedorsallongitudinal
fasciculus.
Projectionstoareasrostraltothehypothalamusarecarriedbythemammillothalamictract,thefornixandterminalstria.
Projectionstoareasofthesympatheticmotorsystem(lateralhornspinalsegmentsT1L2/L3)arecarriedbythehypothalamospinaltractandthey
activatethesympatheticmotorpathway.
Sexualdimorphism
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Severalhypothalamicnucleiaresexuallydimorphici.e.,therearecleardifferencesinbothstructureandfunctionbetweenmalesandfemales.
Somedifferencesareapparenteveningrossneuroanatomy:mostnotableisthesexuallydimorphicnucleuswithinthepreopticarea.Howevermostof
thedifferencesaresubtlechangesintheconnectivityandchemicalsensitivityofparticularsetsofneurons.
Theimportanceofthesechangescanberecognisedbyfunctionaldifferencesbetweenmalesandfemales.Forinstance,malesofmostspeciespreferthe
odorandappearanceoffemalesovermales,whichisinstrumentalinstimulatingmalesexualbehavior.Ifthesexuallydimorphicnucleusislesioned,this
preferenceforfemalesbymalesdiminishes.Also,thepatternofsecretionofgrowthhormoneissexuallydimorphic,andthisisonereasonwhyinmany
species,adultmalesaremuchlargerthanfemales.
Responsivenesstoovariansteroids
Otherstrikingfunctionaldimorphismsareinthebehavioralresponsestoovariansteroidsoftheadult.Malesandfemalesrespondtoovariansteroidsin
differentways,partlybecausetheexpressionofestrogensensitiveneuronsinthehypothalamusissexuallydimorphici.e.,estrogenreceptorsare
expressedindifferentsetsofneurons.
Estrogenandprogesteronecaninfluencegeneexpressioninparticularneuronsorinducechangesincellmembranepotentialandkinaseactivation,
leadingtodiversenongenomiccellularfunctions.Estrogenandprogesteronebindtotheircognatenuclearhormonereceptors,whichtranslocatetothe
cellnucleusandinteractwithregionsofDNAknownashormoneresponseelements(HREs)orgettetheredtoanothertranscriptionfactor'sbindingsite.
Estrogenreceptor(ER)hasbeenshowntotransactivateothertranscriptionfactorsinthismanner,despitetheabsenceofanestrogenresponseelement
(ERE)intheproximalpromoterregionofthegene.Ingeneral,ERsandprogesteronereceptors(PRs)aregeneactivators,withincreasedmRNAand
subsequentproteinsynthesisfollowinghormoneexposure.
Maleandfemalebrainsdifferinthedistributionofestrogenreceptors,andthisdifferenceisanirreversibleconsequenceofneonatalsteroidexposure.
Estrogenreceptors(andprogesteronereceptors)arefoundmainlyinneuronsintheanteriorandmediobasalhypothalamus,notably:
thepreopticarea(whereLHRHneuronsarelocated)
theperiventricularnucleus(wheresomatostatinneuronsarelocated)
theventromedialhypothalamus(whichisimportantforsexualbehavior).
Development
Inneonatallife,gonadalsteroidsinfluencethedevelopmentoftheneuroendocrinehypothalamus.Forinstance,theydeterminetheabilityoffemalesto
exhibitanormalreproductivecycle,andofmalesandfemalestodisplayappropriatereproductivebehaviorsinadultlife.
Ifafemaleratisinjectedoncewithtestosteroneinthefirstfewdaysofpostnatallife(duringthe"criticalperiod"ofsexsteroidinfluence),the
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hypothalamusisirreversiblymasculinizedtheadultratwillbeincapableofgeneratinganLHsurgein
responsetoestrogen(acharacteristicoffemales),butwillbecapableofexhibitingmalesexualbehaviors
(mountingasexuallyreceptivefemale).
Bycontrast,amaleratcastratedjustafterbirthwillbefeminized,andtheadultwillshowfemalesexual
behaviorinresponsetoestrogen(sexualreceptivity,lordosisbehavior).
Inprimates,thedevelopmentalinfluenceofandrogensislessclear,andtheconsequencesarelessunderstood.
Withinthebrain,testosteroneisaromatizedto(estradiol),whichistheprincipalactivehormonefor
developmentalinfluences.Thehumantestissecreteshighlevelsoftestosteronefromaboutweek8offetallife
until56monthsafterbirth(asimilarperinatalsurgeintestosteroneisobservedinmanyspecies),aprocessthat
appearstounderliethemalephenotype.Estrogenfromthematernalcirculationisrelativelyineffective,partly
becauseofthehighcirculatinglevelsofsteroidbindingproteinsinpregnancy.
Sexsteroidsarenottheonlyimportantinfluencesuponhypothalamicdevelopmentinparticular,prepubertal
stressinearlylife(ofrats)determinesthecapacityoftheadulthypothalamustorespondtoanacutestressor.[10]
Unlikegonadalsteroidreceptors,glucocorticoidreceptorsareverywidespreadthroughoutthebraininthe
paraventricularnucleus,theymediatenegativefeedbackcontrolofCRFsynthesisandsecretion,butelsewhere
theirroleisnotwellunderstood.
Mediansagittalsectionofbrainof
humanembryoofthreemonths
Function
Hormonerelease
Thehypothalamushasacentralneuroendocrinefunction,mostnotablybyitscontroloftheanteriorpituitary,
whichinturnregulatesvariousendocrineglandsandorgans.Releasinghormones(alsocalledreleasingfactors)
areproducedinhypothalamicnucleithentransportedalongaxonstoeitherthemedianeminenceortheposterior
pituitary,wheretheyarestoredandreleasedasneeded.[11]
Anteriorpituitary
Inthehypothalamicadenohypophysealaxis,releasinghormones,alsoknownashypophysiotropicor
hypothalamichormones,arereleasedfromthemedianeminence,aprolongationofthehypothalamus,intothe
hypophysealportalsystem,whichcarriesthemtotheanteriorpituitarywheretheyexerttheirregulatory
functionsonthesecretionofadenohypophysealhormones.[12]
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Secretedhormone
Abbreviation
Producedby
Effect
Parvocellular
Thyrotropinreleasinghormone TRH,TRF,or neurosecretorycellsof
(Prolactinreleasinghormone) PRH
theparaventricular
nucleus
Stimulatethyroidstimulatinghormone(TSH)releasefromanterior
pituitary(primarily)
Stimulateprolactinreleasefromanteriorpituitary
Parvocellular
neurosecretorycellsof
Corticotropinreleasinghormone CRHorCRF
theparaventricular
nucleus
Stimulateadrenocorticotropichormone(ACTH)releasefromanterior
pituitary
Dopamine
(Prolactininhibitinghormone)
Growthhormonereleasing
hormone
DAorPIH
Dopamineneuronsof
thearcuatenucleus
GHRH
Neuroendocrineneurons
Stimulategrowthhormone(GH)releasefromanteriorpituitary
oftheArcuatenucleus
GnRHor
Gonadotropinreleasinghormone
LHRH
Somatostatin[13]
(growthhormoneinhibiting
hormone)
Inhibitprolactinreleasefromanteriorpituitary
Stimulatefolliclestimulatinghormone(FSH)releasefromanterior
Neuroendocrinecellsof
pituitary
thePreopticarea
Stimulateluteinizinghormone(LH)releasefromanteriorpituitary
Neuroendocrinecellsof Inhibitgrowthhormone(GH)releasefromanteriorpituitary
SS,GHIH,or
thePeriventricular
Inhibit(moderately)thyroidstimulatinghormone(TSH)releasefrom
SRIF
nucleus
anteriorpituitary
Otherhormonessecretedfromthemedianeminenceincludevasopressin,oxytocin,andneurotensin.[14][15][16][17]
Posteriorpituitary
Inthehypothalamicneurohypophysealaxis,neurohypophysialhormonesarereleasedfromtheposteriorpituitary,whichisactuallyaprolongationofthe
hypothalamus,intothecirculation.
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Secretedhormone
Abbreviation
Producedby
Effect
Oxytocin
Magnocellular
neurosecretorycellsof
OXYorOXT theparaventricular
nucleusandsupraoptic
nucleus
Uterinecontraction
Lactation(letdownreflex)
Vasopressin
(antidiuretichormone)
Magnocellularand
parvocellular
neurosecretorycellsof
ADHorAVP theparaventricular
nucleus,magnocellular
cellsinsupraoptic
nucleus
Increaseinthepermeabilitytowaterofthecellsofdistaltubuleand
collectingductinthekidneyandthusallowswaterreabsorptionand
excretionofconcentratedurine
Itisalsoknownthathypothalamicpituitaryadrenalaxis(HPA)hormonesarerelatedtocertainskindiseasesandskinhomeostasis.Thereisevidence
linkinghyperactivityofHPAhormonestostressrelatedskindiseasesandskintumors.[18]
Stimulation
Thehypothalamuscoordinatesmanyhormonalandbehaviouralcircadianrhythms,complexpatternsofneuroendocrineoutputs,complexhomeostatic
mechanisms,andimportantbehaviours.Thehypothalamusmust,therefore,respondtomanydifferentsignals,someofwhichgeneratedexternallyand
someinternally.DeltawavesignallingarisingeitherinthethalamusorinthecortexinfluencesthesecretionofreleasinghormonesGHRHandprolactin
arestimulatedwhilstTRHisinhibited.
Thehypothalamusisresponsiveto:
Light:daylengthandphotoperiodforregulatingcircadianandseasonalrhythms
Olfactorystimuli,includingpheromones
Steroids,includinggonadalsteroidsandcorticosteroids
Neurallytransmittedinformationarisinginparticularfromtheheart,thestomach,andthereproductivetract
Autonomicinputs
Bloodbornestimuli,includingleptin,ghrelin,angiotensin,insulin,pituitaryhormones,cytokines,plasmaconcentrationsofglucoseand
osmolarityetc.
Stress
Invadingmicroorganismsbyincreasingbodytemperature,resettingthebody'sthermostatupward.
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Olfactorystimuli
Olfactorystimuliareimportantforsexualreproductionandneuroendocrinefunctioninmanyspecies.Forinstanceifapregnantmouseisexposedtothe
urineofa'strange'maleduringacriticalperiodaftercoitusthenthepregnancyfails(theBruceeffect).Thus,duringcoitus,afemalemouseformsa
precise'olfactorymemory'ofherpartnerthatpersistsforseveraldays.Pheromonalcuesaidsynchronisationofoestrusinmanyspeciesinwomen,
synchronisedmenstruationmayalsoarisefrompheromonalcues,althoughtheroleofpheromonesinhumansisdisputed.
Bloodbornestimuli
Peptidehormoneshaveimportantinfluencesuponthehypothalamus,andtodosotheymustpassthroughthebloodbrainbarrier.Thehypothalamusis
boundedinpartbyspecializedbrainregionsthatlackaneffectivebloodbrainbarrierthecapillaryendotheliumatthesesitesisfenestratedtoallowfree
passageofevenlargeproteinsandothermolecules.Someofthesesitesarethesitesofneurosecretiontheneurohypophysisandthemedianeminence.
However,othersaresitesatwhichthebrainsamplesthecompositionoftheblood.Twoofthesesites,theSFO(subfornicalorgan)andtheOVLT
(organumvasculosumofthelaminaterminalis)aresocalledcircumventricularorgans,whereneuronsareinintimatecontactwithbothbloodandCSF.
Thesestructuresaredenselyvascularized,andcontainosmoreceptiveandsodiumreceptiveneuronsthatcontroldrinking,vasopressinrelease,sodium
excretion,andsodiumappetite.Theyalsocontainneuronswithreceptorsforangiotensin,atrialnatriureticfactor,endothelinandrelaxin,eachofwhich
importantintheregulationoffluidandelectrolytebalance.NeuronsintheOVLTandSFOprojecttothesupraopticnucleusandparaventricularnucleus,
andalsotopreoptichypothalamicareas.ThecircumventricularorgansmayalsobethesiteofactionofinterleukinstoelicitbothfeverandACTH
secretion,viaeffectsonparaventricularneurons.
Itisnotclearhowallpeptidesthatinfluencehypothalamicactivitygainthenecessaryaccess.Inthecaseofprolactinandleptin,thereisevidenceof
activeuptakeatthechoroidplexusfromthebloodintothecerebrospinalfluid(CSF).Somepituitaryhormoneshaveanegativefeedbackinfluenceupon
hypothalamicsecretionforexample,growthhormonefeedsbackonthehypothalamus,buthowitentersthebrainisnotclear.Thereisalsoevidencefor
centralactionsofprolactin.
Findingshavesuggestedthatthyroidhormone(T4)istakenupbythehypothalamicglialcellsintheinfundibularnucleus/medianeminence,andthatit
ishereconvertedintoT3bythetype2deiodinase(D2).Subsequenttothis,T3istransportedintothethyrotropinreleasinghormone(TRH)producing
neuronsintheparaventricularnucleus.Thyroidhormonereceptorshavebeenfoundintheseneurons,indicatingthattheyareindeedsensitivetoT3
stimuli.Inaddition,theseneuronsexpressedMCT8,athyroidhormonetransporter,supportingthetheorythatT3istransportedintothem.T3couldthen
bindtothethyroidhormonereceptorintheseneuronsandaffecttheproductionofthyrotropinreleasinghormone,therebyregulatingthyroidhormone
production.[19]
Thehypothalamusfunctionsasatypeofthermostatforthebody.[20]Itsetsadesiredbodytemperature,andstimulateseitherheatproductionand
retentiontoraisethebloodtemperaturetoahighersettingorsweatingandvasodilationtocoolthebloodtoalowertemperature.Allfeversresultfroma
raisedsettinginthehypothalamuselevatedbodytemperaturesduetoanyothercauseareclassifiedashyperthermia.[20]Rarely,directdamagetothe
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hypothalamus,suchasfromastroke,willcauseafeverthisissometimescalledahypothalamicfever.However,itismorecommonforsuchdamageto
causeabnormallylowbodytemperatures.[20]
Steroids
Thehypothalamuscontainsneuronsthatreactstronglytosteroidsandglucocorticoids(thesteroidhormonesoftheadrenalgland,releasedinresponse
toACTH).Italsocontainsspecializedglucosesensitiveneurons(inthearcuatenucleusandventromedialhypothalamus),whichareimportantfor
appetite.ThepreopticareacontainsthermosensitiveneuronstheseareimportantforTRHsecretion.
Neural
Oxytocinsecretioninresponsetosucklingorvaginocervicalstimulationismediatedbysomeofthesepathwaysvasopressinsecretioninresponseto
cardiovascularstimuliarisingfromchemoreceptorsinthecarotidbodyandaorticarch,andfromlowpressureatrialvolumereceptors,ismediatedby
others.Intherat,stimulationofthevaginaalsocausesprolactinsecretion,andthisresultsinpseudopregnancyfollowinganinfertilemating.Inthe
rabbit,coituselicitsreflexovulation.Inthesheep,cervicalstimulationinthepresenceofhighlevelsofestrogencaninducematernalbehaviorinavirgin
ewe.Theseeffectsareallmediatedbythehypothalamus,andtheinformationiscarriedmainlybyspinalpathwaysthatrelayinthebrainstem.
StimulationofthenipplesstimulatesreleaseofoxytocinandprolactinandsuppressesthereleaseofLHandFSH.
Cardiovascularstimuliarecarriedbythevagusnerve.Thevagusalsoconveysavarietyofvisceralinformation,includingforinstancesignalsarising
fromgastricdistensionoremptying,tosuppressorpromotefeeding,bysignallingthereleaseofleptinorgastrin,respectively.Againthisinformation
reachesthehypothalamusviarelaysinthebrainstem.
Inadditionhypothalamicfunctionisresponsivetoandregulatedbylevelsofallthreeclassicalmonoamineneurotransmitters,noradrenaline,
dopamine,andserotonin(5hydroxytryptamine),inthosetractsfromwhichitreceivesinnervation.Forexample,noradrenergicinputsarisingfromthe
locuscoeruleushaveimportantregulatoryeffectsuponcorticotropinreleasinghormone(CRH)levels.
Controloffoodintake
Theextremelateralpartoftheventromedialnucleusofthehypothalamusisresponsibleforthecontroloffoodintake.Stimulationofthisareacauses
increasedfoodintake.Bilaterallesionofthisareacausescompletecessationoffoodintake.Medialpartsofthenucleushaveacontrollingeffectonthe
lateralpart.Bilaterallesionofthemedialpartoftheventromedialnucleuscauseshyperphagiaandobesityoftheanimal.Furtherlesionofthelateralpart
oftheventromedialnucleusinthesameanimalproducescompletecessationoffoodintake.
Therearedifferenthypothesesrelatedtothisregulation:[22]
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1.Lipostatichypothesis:Thishypothesisholdsthatadiposetissueproducesahumoral
signalthatisproportionatetotheamountoffatandactsonthehypothalamusto
decreasefoodintakeandincreaseenergyoutput.Ithasbeenevidentthatahormone
leptinactsonthehypothalamustodecreasefoodintakeandincreaseenergyoutput.
2.Gutpeptidehypothesis:gastrointestinalhormoneslikeGrp,glucagons,CCKand
othersclaimedtoinhibitfoodintake.Thefoodenteringthegastrointestinaltract
triggersthereleaseofthesehormones,whichactonthebraintoproducesatiety.The
braincontainsbothCCKAandCCKBreceptors.
3.Glucostatichypothesis:Theactivityofthesatietycenterintheventromedialnucleiis
probablygovernedbytheglucoseutilizationintheneurons.Ithasbeenpostulated
thatwhentheirglucoseutilizationislowandconsequentlywhenthearteriovenous
bloodglucosedifferenceacrossthemislow,theactivityacrosstheneuronsdecrease.
Undertheseconditions,theactivityofthefeedingcenterisuncheckedandthe
individualfeelshungry.Foodintakeisrapidlyincreasedbyintraventricular
administrationof2deoxyglucosethereforedecreasingglucoseutilizationincells.
4.Thermostatichypothesis:Accordingtothishypothesis,adecreaseinbody
temperaturebelowagivensetpointstimulatesappetite,whereasanincreaseabove
thesetpointinhibitsappetite.
Peptidehormonesandneuropeptidesthatregulate
feeding[21]
Peptidesthatincrease
Peptidesthatdecrease
feedingbehavior
feedingbehavior
Ghrelin
Leptin
NeuropeptideY
(,,)Melanocyte
stimulatinghormones
Agoutirelatedpeptide
Cocaineandamphetamine
regulatedtranscriptpeptides
Orexins(A,B)
Corticotropinreleasing
hormone
Melaninconcentrating
hormone
Cholecystokinin
Galanin
Insulin
Glucagonlikepeptide1
Fearprocessing
Themedialzoneofhypothalamusispartofacircuitrythatcontrolsmotivatedbehaviors,likedefensivebehaviors.[23]AnalysesofFoslabelingshowed
thataseriesofnucleiinthe"behavioralcontrolcolumn"isimportantinregulatingtheexpressionofinnateandconditioneddefensivebehaviors.[24]
Antipredatorydefensivebehavior
Exposuretoapredator(suchasacat)elicitsdefensivebehaviorsinlaboratoryrodents,evenwhentheanimalhasneverbeenexposedtoacat.[25]Inthe
hypothalamus,thisexposurecausesanincreaseinFoslabeledcellsintheanteriorhypothalamicnucleus,thedorsomedialpartoftheventromedial
nucleus,andintheventrolateralpartofthepremammillarynucleus(PMDvl).[26]Thepremammillarynucleushasanimportantroleinexpressionof
defensivebehaviorstowardsapredator,sincelesionsinthisnucleusabolishdefensivebehaviors,likefreezingandflight.[26][27]ThePMDdoesnot
modulatedefensivebehaviorinothersituations,aslesionsofthisnucleushadminimaleffectsonpostshockfreezingscores.[27]ThePMDhasimportant
connectionstothedorsalperiaqueductalgray,animportantstructureinfearexpression.[28][29]Inaddition,animalsdisplayriskassessmentbehaviorsto
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theenvironmentpreviouslyassociatedwiththecat.FoslabeledcellanalysisshowedthatthePMDvlisthemostactivatedstructureinthehypothalamus,
andinactivationwithmuscimolpriortoexposuretothecontextabolishesthedefensivebehavior.[26]Therefore,thehypothalamus,mainlythePMDvl,
hasanimportantroleinexpressionofinnateandconditioneddefensivebehaviorstoapredator.
Socialdefeat
Likewise,thehypothalamushasaroleinsocialdefeat:Nucleiinmedialzonearealsomobilizedduringanencounterwithanaggressiveconspecific.
ThedefeatedanimalhasanincreaseinFoslevelsinsexuallydimorphicstructures,suchasthemedialpreopticnucleus,theventrolateralpartof
ventromedialnucleus,andtheventralpremammilarynucleus.[30]Suchstructuresareimportantinothersocialbehaviors,suchassexualandaggressive
behaviors.Moreover,thepremammillarynucleusalsoismobilized,thedorsomedialpartbutnottheventrolateralpart.[30]Lesionsinthisnucleusabolish
passivedefensivebehavior,likefreezingandthe"ontheback"posture.[30]
Sexualorientation
AccordingtoD.F.Swaab,writinginaJuly2008paper,"Neurobiologicalresearchrelatedtosexualorientationinhumansisonlyjustgathering
momentum,buttheevidencealreadyshowsthathumanshaveavastarrayofbraindifferences,notonlyinrelationtogender,butalsoinrelationto
sexualorientation."[31]
Swaabfirstreportedontherelationshipbetweensexualorientationinmalesandthehypothalamus's"clock",thesuprachiasmaticnucleus(SCN).In
1990,SwaabandHofman[32]reportedthatthesuprachiasmaticnucleusinhomosexualmenwassignificantlylargerthaninheterosexualmen.Thenin
1995,Swaabetal.[33]linkedbraindevelopmenttosexualorientationbytreatingmaleratsbothpreandpostnatallywithATD,anaromataseblockerin
thebrain.ThisproducedanenlargedSCNandbisexualbehaviorintheadultmalerats.In1991,LeVayshowedthatpartofthesexuallydimorphic
nucleus(SDN)knownasthe3rdinterstitialnucleusoftheanteriorhypothalamus(INAH3),isnearlytwiceaslarge(intermsofvolume)inheterosexual
menthaninhomosexualmenandheterosexualwomen.However,astudyin1992hasshownthatthesexuallydimorphnucleusofthepreopticarea,
whichincludetheINAH3,areofsimilarsizeinhomosexualmaleswhodiedofAIDStoheterosexualmales,andthereforelargerthanfemale.This
clearlycontradictsthehypothesisthathomosexualmaleshaveafemalehypothalamus.Furthermore,theSCNofhomosexualmalesisextremelylarge
(boththevolumeandthenumberofneuronsaretwiceasmanyasinheterosexualmales).Theseareasofthehypothalamushavenotyetbeenexploredin
homosexualfemalesnorbisexualmalesnorfemales.Althoughthefunctionalimplicationsofsuchfindingsstillhaven'tbeenexaminedindetail,they
castseriousdoubtoverthewidelyacceptedDrnerhypothesisthathomosexualmaleshavea"femalehypothalamus"andthatthekeymechanismof
differentiatingthe"malebrainfromoriginallyfemalebrain"istheepigeneticinfluenceoftestosteroneduringprenataldevelopment.[34][35]
In2004and2006,twostudiesbyBerglund,Lindstrm,andSavic[36][37]usedpositronemissiontomography(PET)toobservehowthehypothalamus
respondstosmellingcommonodors,thescentoftestosteronefoundinmalesweat,andthescentofestrogenfoundinfemaleurine.Thesestudies
showedthatthehypothalamusofheterosexualmenandhomosexualwomenbothrespondtoestrogen.Also,thehypothalamusofhomosexualmenand
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