Kutyák Kontextusfüggő Agresszív Viselkedésének Viselkedésélettani Jellemzése Doktori Értekezés

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Kutyák Kontextusfüggő Agresszív Viselkedésének Viselkedésélettani Jellemzése Doktori Értekezés Eötvös Loránd Tudományegyetem Biológia Doktori Iskola Vezető: Prof. Dr. Erdei Anna Etológia Doktori Program Vezető: Dr. Miklósi Ádám Igyártó (Horváth) Zsuzsánna Kutyák kontextusfüggő agresszív viselkedésének viselkedésélettani jellemzése Doktori értekezés Témavezető: Dr. Miklósi Ádám egyetemi docens Eötvös Loránd Tudományegyetem Etológia Tanszék 2009 ”A stressz az élet sava-borsa. A stresszmentes állapot a halál.” ~Selye János~ ”A felfedezés lényege: látni azt, amit már mindenki látott, de olyat gondolni, amit senki más nem gondolt róla.” ~Szent-Györgyi Albert~ 2 TARTALOMJEGYZÉK 1. Bevezetés......................................................................................................................5 1.1. Agresszív viselkedésformák.......................................................................................6 1.1.1. Az agresszivitás funkcionális magyarázata...............................................................6 1.1.2. Az agresszív viselkedés élettani tényezői..................................................................9 1.1.2.1. Hormonok...............................................................................................................9 1.1.2.2. Neurotranszmitterek..............................................................................................12 1.1.2.3. Neurális struktúrák...............................................................................................14 1.1.2.4. Genetikai háttér....................................................................................................14 1.2. A kutya agresszív viselkedésformái........................................................................16 1.2.1. A kutyák agresszív viselkedését befolyásoló biológiai tényezők............................18 1.2.2. Az agresszív viselkedés megfigyelése kutyáknál....................................................20 1.3. Stressz........................................................................................................................23 1.3.1. A stresszről általánosan............................................................................................23 1.3.1.1. Az akut és krónikus stressz....................................................................................25 1.3.1.2 A stresszre adott választ befolyásoló tényezők.......................................................26 1.3.2. A kortizol szerepe a stresszválasz kialakulásában...................................................27 1.4. A kutya stresszviselkedésének aktiválása és a kortizol jelző szerepe...................28 1.5. Agresszivitás és stressz.............................................................................................30 1.5.1. A győzelem és vereség hatása a tesztoszteron szintre.............................................32 1.5.2. Megküzdési stílusok megjelenése stresszhelyzetben...............................................33 1.6. A kutya és ember közötti szociális kapcsolat..........................................................35 1.6.1. Interspecifikus szociális hatások a belső állapotra: Az ember hatása a kutyára......38 1.6.2. Interspecifikus szociális hatások a belső állapotra: A kutya hatása az emberre......39 2. Célkitűzések.............................................................................................................41 2.1. Szeparáció..................................................................................................................43 2.2. Fenyegető megközelítés............................................................................................44 2.3. Játékos helyzet...........................................................................................................45 2.4. Vizsgaszituáció..........................................................................................................46 3. Módszertani problémák.......................................................................................48 3.1. A kutyák kortizolszintjét befolyásoló tényezők......................................................48 3.2. A kortizolszint nyálból történő mérésének jogosultsága és előnyei......................49 3.3. Kortizol mérése nyálból............................................................................................50 4. Vizsgálatok................................................................................................................53 4.1. Általános módszer.....................................................................................................53 4.1.1. Alanyok....................................................................................................................53 4.1.2. Időpont és helyszín..................................................................................................54 4.1.3. A nyál kortizol és tesztoszteron koncentrációjának mérése....................................54 4.2. Szeparáció..................................................................................................................55 4.2.1. Módszer....................................................................................................................55 4.2.2. Eredmények.............................................................................................................59 4.2.3. Eredmények értékelése............................................................................................65 3 4.3. Fenyegető megközelítés............................................................................................67 4.3.1. Módszer....................................................................................................................67 4.3.2. Eredmények.............................................................................................................72 4.3.3. Eredmények értékelése............................................................................................79 4.4. Játékos helyzet...........................................................................................................81 4.4.1. Módszer....................................................................................................................81 4.4.2. Eredmények.............................................................................................................86 4.4.3. Eredmények összefoglalása.....................................................................................91 4.5. Vizsgaszituáció..........................................................................................................93 4.5.1. Módszer....................................................................................................................93 4.5.2. Eredmények.............................................................................................................99 4.5.3. Eredmények összefoglalása...................................................................................105 5. Diszkusszió..............................................................................................................108 5.1. Stressz vizsgálata kutyán, avagy tesztjeink jogosultsága....................................108 5.2. A szeparációs stressz megjelenése rendőrkutyák esetében.................................109 5.3. Agresszivitás és stressz összefüggésének vizsgálata kutyán: Különböző megküzdési stílusok kimutatása szociális stressz esetén.....................................113 5.4. Az ember viselkedésének befolyásoló hatása a kutyára egy játékos szociális interakció esetén.....................................................................................................115 5.5. Kompenzációs hipotézis..........................................................................................118 5.6. A kor és munka típusa, mint stresszválaszt befolyásoló tényezők......................122 6. Konklúzió................................................................................................................124 Köszönetnyilvánítás.................................................................................................125 Irodalomjegyzék........................................................................................................127 Összefoglalás...............................................................................................................149 Summary......................................................................................................................150 4 1. BEVEZETÉS Az agresszivitás és stressz közötti szoros kapcsolat ma már általánosan ismert és elfogadott tény. A szociális agresszív interakciók stresszelőek, és a stressz következtében felszabaduló glükokortikoidok (stresszhormonok) serkenthetik vagy gátolhatják az agresszivitást a hormonhatás időtartamától függően. Akut stressz esetén, amikor az állat még képes megküzdeni az őt ért kihívással, a termelődő glükokortikoidok hozzájárulnak az agresszív viselkedés teljes kifejeződéséhez. Ezzel szemben krónikus stressz esetén, amikor az állat már hosszabb ideje képtelen megszabadulni a stresszt okozó tényezőtől, a glükokortikoidok krónikus túltermelése gátolja az agresszivitást. Ezek alapján, a rövid ideig tartó stressz aktív agresszív válasszal ellensúlyozódik, ugyanakkor krónikus, kontrollálhatatlan és megjósolhatatlan körülmények között a túlélés elősegítése, a kockázatok és energiaveszteségek kerülése az előnyös. Az agresszivitás és stressz együttes jelenlétét elsősorban falkában vagy csoportban élő állatok esetében tanulmányozták, ahol a rangsor ugyan lehetővé
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