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倉鼠也行社會性訊息襲取且會引發觀察學習當下以及事後的改變

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୯ҥᆵ᡼εᏢ౛ᏢଣЈ౛Ꮲࣴز܌

ᅺγፕЎ

Graduate Institute of Psychology College of Science

National Taiwan University Master Thesis

ঊႵΨՉޗ཮܄ૻ৲᠍ڗЪ཮ЇวᢀჸᏢಞ྽Π!

аϷ٣ࡕޑׯᡂ !

Hamsters Eavesdrop Too and The Consequent Changes during and after Observational Learning

೾ܴے Ming-Tsung Kuo

ࡰᏤ௲௤ǺᒘЎ஡ റγ Advisor: Wen-Sung Lai, Ph.D.

ύ๮҇୯ 98 ԃ 7 Д

July, 2009

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ᇞᖴ

ӳǼᕴᆉֹԋ೭ҁፕЎΑǴ೾ܴےǴ༸ளӳǼӃ๏ԾρٰঁඓᖂփǴ୙Ǽ୙Ǽ

୙Ǽ୙ǼΟԃΑǴӣགྷ྽߃Ǵϙሶ೿όᔉǴ൩೭ኬ஥๱ᅈ๚዗Ո፯Ε೭ঁࣽᏢᘀ

݅ǴΟԃၸѐΑǴᗨฅၹᡏᡒ໾ǴՠЈǴ׳уय֧ǶाགᖴޑΓϼӭǴόѝࢂᖴ ϺǶགᖴךޑР҆ᆶঢঢکۆۆǴؒԖգॺǴό཮ԖϞϺޑךᆶךޑܴϺǶךޑ

ࡰᏤ௲௤ǴᒘЎ஡ԴৣǴᖴᖴா೭Οԃٰޑаيբ߾ᆶ໼᠋ǴᖴᖴǼགᖴలζԶ ۆǴךࣴز܌ғఱύޑ຦ΓǴᖴᖴா΋ޔаٰעך྽ٽη૓ྣ៝ǴᖴᖴǶఉٍ۪

ԴৣǴᖴᖴாǴӵ݀ϖԃ߻ؒԖ᠋ၸா΢ளፐǴךёૈᗋόᏢคೌǴόޕՖᒏғ

౛Ј౛ᏢǴ׳ᖴᖴாѳВޑᜢЈǶس΢ޑߋӃғǴᖴᖴாதӧךႝတрୢᚒਔр Ћ࣬շǴϼགᖴΑǼᖴᖴᗛቺᏦԴৣᆶߋඵ፣Դৣӧךޑჴᡍ΢๏ϒڐշᆶࡰ

ᏤǴᗨฅ࣬ೀਔ໔όߏǴՠߚதᖴᖴǶᖴᖴ৪჏ֻԴৣᆶᎄ໡ᖚԴৣᜫཀ྽ךޑ α၂ہ঩٠๏ϒᝊ຦ޑࡌ᝼Ƕғ౛܌ޑࡼ࣓࣓Ꮲۆᆶε໵Ꮲ׌Ǵᖴᖴգॺӧךޑ ჴᡍ΢๏ϒڐշᆶࡰᏤǴᖴᖴǼሺོǵৎྍǵЎᇬǵࣁฺǵӵదǵаϷॕ⪭Ǵག ᖴգॺჴᡍ΢ޑڐշǶΓيፏ૓ԵᡍǴόࢂाᔐॹգ Զࢂाԋ൩գǴࠂᏌᏢߏǵ

܃ЎǵᝬЬǵаϷ㾈ሺǴғڮύԖգॺޑр౜ࢂךࣴز܌ޑВη္Ǵԋߏ΢ࡐε ޑ୏ΚٰྍǴᖴᖴǼШၲᏢߏǵ࠮ើᏢۆǵаϷไ޼ᏢۆǴᖴᖴգॺӧךԖሡा

ਔεΚ࣬շǶζΐᓓ᡺ޑӚՏߓ࠰ॺǴߚதགᖴգॺ೭Οԃჹךޑྣ៝Ǵத๏ך у๼Ǵϼགᖴ୚Ǽགᖴך܌Ԗޑӳ϶ॺǴᖴᖴգॺૈх৒ךǴ҂ٰᗋፎӭӭх֖Ƕ നࡕǴךाߚதᖴᖴமԶԖΚམӧךުޑออǴךന഻៿یΑǼЋ౐ЋǴ΋ଆ຾

؁փǶ

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(4)

ᄔा

ᢀჸᏢಞǴཀࡰғЋ೸ၸᢀჸдΓޑՉ୏຾ԶಞளཥޑՉࣁ܈ૻ৲Ǵ೭ᅿᏢ ಞБԄςӧόӕނᅿޑፏӭՉࣁύ೏᛾ჴၸǶ೸ၸᢀჸдΓޑޗ཮ϕ୏຾Զவύ

๧ڗрૻ৲ޑ೭ᅿᏢಞБԄ೏ᆀࣁޗ཮܄ૻ৲᠍ڗǹၸѐ࣬ᜢޑࣴزεӭଞჹΓ ᜪаϷࢌ٤ങᜪᆶചᜪǴՠଞჹΓᜪаѦޑথ٢ᜪ୏ނᗋ҂මԖၸǶճҔ௶ճ٥

໢܄ঊႵڀԖϐሦୱװᔐՉࣁǴךॺว৖р΋ঁՉࣁᏹբБݤٰ௖૸ޗ཮܄ૻ৲

᠍ڗǶҁፕЎύӅԖΟঁЬჴᡍǴҞޑӧ௖૸ᢀჸޣೱុΟϺჹٿঁҢጄޣޑѺ ର܈ύ܄٣ҹ຾Չᢀჸਔ܌ЇวᢀჸᏢಞ྽ΠаϷ٣ࡕϐՉࣁᆶᓸΚລᅟᆾޑ ׯᡂǶӧჴᡍ΋ύǴ҂ԖѺର࿶ᡍޑ໢܄ᢀჸޣǴӧΟϺᢀჸ่״ࡕҥջ܈ࢂ΋

ϺࡕǴܭ U ׎଎ࢭϣ཮זೲޑᖿ߈٠ᡉ๱Ӧ޸ၨӭਔ໔ѐ௖઩គৎҢጄޣǴᡉ Ң໢܄ᢀჸޣКၨԖᑫ፪ᆶគৎҢጄޣ຾Չϕ୏Ƕ࣬ၨܭჴᡍ΋ύ٬ҔΑ҂ԖѺ ର࿶ᡍϐᢀჸޣǴӧჴᡍΒύ߾ࢂ٬ҔΑ຾ՉᢀჸᏢಞ߻΋Ϻමᎁၶढ़ғװᔐޣ

਋௳ၸ΋ԛޑ໢܄ᢀჸޣǴ٠ว౜೭٤ᢀჸޣӧ࿶ᐕೱុΟϺᢀჸѺର٣ҹޑ྽

ΠаϷࡕុӧ U ׎଎ࢭύޑҥջکߏයෳ၂΢Ǵ֡཮߄౜рᆶ߻΋ঁჴᡍֹӄ

࣬ϸޑՉࣁኳԄȐх֖Ǻၨӭޑᕨߡ௨ܫǵၨӭޑ଒າᆶ଑ᗉǵаϷၨϿޑ௖઩

ਔ໔ȑǴՠԜਏ݀٠ό཮р౜ӧೱុΟϺᢀჸύ܄٣ҹϐԖ਋௳࿶ᡍޑᢀჸޣ

΢Ƕ೭٤่݀ᡉҢΑ਋௳࿶ᡍૈԖਏӦቹៜڙ਋ঊႵځࡕޑՉࣁ߄౜ǶӧჴᡍΟ ύǴ٬ҔΑߚߟΕ܄ޑБݤٰୀෳᢀჸޣᕨߡϣҜ፦ᎇжᖴނȐFCMȑޑ֖ໆǶ

ӧჴᡍΟ A ύǴ೸ၸ؂ 3 ঁλਔᇆ໣΋ԛᕨߡኬҁޑ௦ኬБݤǴࣴز่݀ᡉҢ ӧᐕ࿶΋ԛװᔐϕ୏ࡕǴᒡৎᕨߡϣޑҜ፦ᎇжᖴނᐚࡋܭ٣ҹࡕޑಃΟᆶಃΒ

Μ΋λਔ཮Ԗܴᡉޑ΢ϲǴՠ೭ኬޑׯᡂ٠ό཮วғӧគৎي΢ǶӧჴᡍΟ B ύǴ߾٬ҔΑ٩ᏵჴᡍΟ A ่݀অׯၸޑ؂ 18 λਔᕨߡ௦໣ݤٰϩ݋ჴᡍΒύ ޑᢀჸޣ܌௨ܫрޑᕨߡኬҁǶӧΟϺޑᢀჸύǴԖ਋௳࿶ᡍޑ໢܄ᢀჸޣӧᢀ ჸѺର٣ҹਔǴځ௨ᕨໆԖܴᡉޑቚуǴՠځᕨߡϣҜ፦ᎇжᖴނޑᐚࡋࠅ࣬ჹ

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եܭᢀჸύ܄ϕ୏٣ҹޑᢀჸޣǶӕኬӦӧ่״ᢀჸࡕ΋ϺǴय़ჹគৎҢጄޣ߄

౜ᐟਗ਼ޑᢀჸޣي΢Ψ࡭ុໆෳрᕨߡϣҜ፦ᎇжᖴނफ़եޑ௃׎Ƕᆕӝа΢܌

ॊǴ೭٤่݀ᡉҢΑ໢܄ঊႵΨૈ຾Չޗ཮܄ૻ৲᠍ڗǴЪঁᡏၸѐޗҬ਋௳ޑ

࿶ᡍ཮ᡉ๱Ӧቹៜځࡕុၶډ೭٤ወӧჹЋਔૻ৲ޑ٬ҔаϷຑ՗ޑБԄǶᙖҗ ᢀჸ܌ᕇளٰޑૻ৲ǴόѝቹៜΑғ౛΢ޑࡡۓ܄ǴΨቹៜ๱ᒿࡕޑՉࣁϸᔈǶ

ᣂ᝶ڗΚᨠኘᖂ฾Εषᄎࢤಛஒ᦭࠷ΕᚘԺ֘ᚨΕސᚰࢤΕᜈঁփؼᔆᔧז᝔ढΕ ପቕΖ

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Hamsters Eavesdrop Too and The Consequent Changes during and after Observational Learning

Ming-Tsung Kuo

Abstract

Observational learning, refers naïve individual acquires new behavior or information via observation of others’ doing, has been demonstrated in a variety of behaviors in different species. Gathering information from social interactions between others is termed eavesdropping which has been studied mainly in humans and certain kinds of fishes and birds, but not in other mammals. Using male golden hamsters and their agonistic behaviors, we developed a behavioral method to study eavesdropping.

In this thesis, there are 3 experiments aiming at investigating consequential behavioral and hormonal changes in different groups of observers during and after the

observation of two male demonstrators either fighting or encountering neutrally across a 3-day observational learning. In experiment 1, fighting naïve males approach quickly and spent significantly more time investigating the winning demonstrator in the U-maze immediately or one day after the 3-day observation indicating that they are more interested in interacting with the winners rather than the losers. In

experiment 2, in contrast to those males in experiment 1, observers previously received one defeated experience from a novel fighter one day before the

observational learning displayed completely opposite behavioral patterns (i.e., more defecation, more fleeing behavior and avoidance, and less investigation time toward the winning demonstrator) during the observation of the 3-day aggressive interactions and in the following immediate and long-term U-maze tests whereas such effects did

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not appear in the defeated males observed 3-day neutral interactions. These results indicate that the experience of social defeat can effectively influence subsequent behaviors in defeated hamsters. In experiment 3, a non-invasive method was applied to measure the levels of fecal cortisol metabolites (FCM). In experiment 3A, fecal samples were collected every 3 hours and our data indicate that a single fighting experience increased FCM level in the losers 3 hours and 21 hours after the

aggressive encounter but has no effect on the winners at any time point. In experiment 3B, a modified 18-hour fecal collection procedure was adopted to analyze those fecal samples collected from observers in experiment 2. During the 3-day observation, these defeated males that observed aggressive interactions defecated more sequentially but their FCM level was relatively decreased compared to those that observed neutral encounters. One day after the last observation, the resultant decrease of FCM level was remained in those high responders that faced the winning

demonstrators. Taken together, these results suggest that male hamsters eavesdrop too and the defeated experience significantly affects the ways they use the information and evaluate their potential opponents in subsequent encounter. The information gathered during the observation not only sufficiently affects physical homeostasis but also influences sequential behavioral responses.

Key words: observational learning, eavesdropping, stress response, aggression, fecal cortisol metabolites, hamster.

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Ҟԛ!

!

ಃ΋കʳ ᆣፕ////////////////////////////////////////////////////////1!

ಃ΋࿯ʳ Їق////////////////////////////////////////////////////1 ᢀჸᏢಞ////////////////////////////////////////////////////1

ಃΒ࿯ʳ ӣ៝౜ϞӸӧܭ୏ނي΢ޑᢀჸᏢಞࣴز////////////////////3

൘ǵᆶғӸԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////////3 ມǵᆶ܄ᆶᕷ़ԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////6

ୖǵᆶٛᑇԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////////7 စǵኳү////////////////////////////////////////////////////9

ಃΟ࿯ʳ ޗ཮܄ૻ৲᠍ڗ/////////////////////////////////////////12

൘ǵวғܭޗ཮܄ᝡݾύޑޔௗຑ՗ᆶ໔ௗຑ՗/////////////////12 ມǵޗ཮܄ᝡݾύଞჹޗ཮܄ૻ৲᠍ڗӧ୏ނي΢ޑࣴز/////////14

ୖǵঁᡏ࿶ᡍჹޗ཮ᝡݾύޑޗ཮܄ૻ৲᠍ڗޑቹៜ/////////////17

ಃѤ࿯ʳ ҁፕЎޑࣴز᝼ᚒᆶ฼ౣ/////////////////////////////////18

൘ǵࣴز᝼ᚒ///////////////////////////////////////////////18 ມǵਡЈୢᚒᆶှ،฼ౣ/////////////////////////////////////22

ಃΒകʳ ჴᡍ׷਑ᆶБݤ/////////////////////////////////////////////25!

ಃ΋࿯ʳ ჴᡍ΋/////////////////////////////////////////////////25

ಃΒ࿯ʳ ჴᡍΒ/////////////////////////////////////////////////33

ಃΟ࿯ʳ ჴᡍΟ/////////////////////////////////////////////////36

൘ǵჴᡍΟ A///////////////////////////////////////////////37 ມǵჴᡍΟ B///////////////////////////////////////////////41

ಃΟകʳ ่݀ᆶ૸ፕ/////////////////////////////////////////////////45!

ಃ΋࿯ʳ ჴᡍ΋/////////////////////////////////////////////////45

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௖૸҂ԖޗҬ࿶ᡍޑᢀჸޣӧޗ཮܄ૻ৲᠍ڗޑᐕำύᆶࡕុૻ৲٬

ҔਔޑՉࣁ߄౜/////////////////////////////////////////////45

ಃΒ࿯ʳ ჴᡍΒ/////////////////////////////////////////////////52

௖૸ڀԖ΋ԛޗҬ਋௳࿶ᡍޑᢀჸޣӧޗ཮܄ૻ৲᠍ڗޑᐕำύᆶࡕ

ុૻ৲٬ҔਔޑՉࣁ߄౜/////////////////////////////////////52

ಃΟ࿯ʳ ჴᡍΟ A///////////////////////////////////////////////62

௖૸΋ԛޗҬ਋௳࿶ᡍΠޑᓸΚϸᔈ///////////////////////////62

ಃѤ࿯ʳ ჴᡍΟ B///////////////////////////////////////////////67

௖૸ڀԖ΋ԛޗҬ਋௳࿶ᡍޑᢀჸޣӧޗ཮܄ૻ৲᠍ڗޑᐕำύᆶࡕ

ុޑૻ৲٬ҔਔޑᓸΚϸᔈ///////////////////////////////////67

ಃѤകʳ ᆕӝ૸ፕ///////////////////////////////////////////////////73!

ಃ΋࿯ʳ ճҔ໢܄௶ճ٥ঊႵ୏ނኳԄࣴز࣬ᜢ᝼ᚒޑᓬ༈///////////74

ಃΒ࿯ʳ ҂ԖޗҬ࿶ᡍΠޑእЋຑ՗ᐕำ///////////////////////////77

ಃΟ࿯ʳ ڀԖޗҬ਋௳࿶ᡍΠޑእЋຑ՗ᐕำ///////////////////////78

ಃѤ࿯ʳ ঁᡏޗҬ਋௳࿶ᡍჹޗ཮܄ૻ৲᠍ڗᐕำᆶࡕុޑእЋຑ՗ᐕำ ϐቹៜ܄///////////////////////////////////////////////79

ಃϖ࿯ʳ ޗ཮܄ૻ৲ௗڙਔᆶঁᡏӆᇡਔ܌ճҔޑጕ઩///////////////81

ಃϤ࿯ʳ ᓸΚ࿶ᡍΠޑঁᡏৡ౦܄/////////////////////////////////83

ಃΎ࿯ʳ ҁፕЎޑଅ᝘ᆶځ҂ٰޑᔈҔ/////////////////////////////85

ୖԵЎ᝘///////////////////////////////////////////////////////////87!

ߕკᇥܴ///////////////////////////////////////////////////////////95!

ߕკ//////////////////////////////////////////////////////////////103!

ߕ߄//////////////////////////////////////////////////////////////117!

ߕᒵ//////////////////////////////////////////////////////////////127

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ಃ΋കʳ ᆣፕ

ಃ΋࿯ʳ Їق

ȸIt is not the strongest of the species that survive, nor the most intelligent, but the one most responsive to changeȹȋ Charles Darwin

ᏢಞǴջނᅿࣁΑૈ፾ᔈᕉნזೲᡂϯ܌ᄽϯрޑ΋ᅿૈΚǴѬ཮٩ᕉნޑ ᡂϯԶ଺р፾྽ޑፓ᏾Ǵ܌٩ᒘޑջࣁၸѐჹ၀ڈᐟ܈၀ᕉნޑ૶ᏫǶᗨฅӵ ԜǴᏢಞޣ۳۳ሡाᒃԾѐ࿶ᐕ٠бржሽǴωளаᕇள࿶ᡍǴٯӵǺ೸ၸᒃԾ ӞΠࢌނᡏࡕᏤठғੰǴωΑှډԜނᡏࢂόё१ޑǶ

ᢀჸᏢಞ

ঁᡏᏢಞȐindividual learningȑ೯தሡा೸ၸޔௗѐ࿶ᐕ٣ҹωૈᏢಞډǴ Ψ۳۳ӢԜбрΑ΋ۓޑжሽǹ࣬ჹԶقǴќ΋ᅿ໔ௗԄޑᏢಞ߾ࢂ೸ၸᢀჸځ дΓޑ࿶ᡍǴ൩ૈڙډቹៜ܈ᕇளૻ৲Ǵ೭൩ࢂ܌ᒏޑᢀჸᏢಞȐobservational learningȑǶӧঁᡏᏢಞᐕำύǴ೯தሡाᐕ࿶΋ۓำࡋޑ၂ᇤᆶ჋၂Ǵωૈගଯ ಞள၀Չࣁޑёૈ܄ǹ࣬ჹԶقǴᢀჸᏢಞѝሡा೸ၸᢀჸձΓޑՉࣁ߄౜Ǵ൩

ૈගଯಞள၀Չࣁޑёૈ܄ǴӢԜᢀჸᏢಞёаεεफ़ե܌ሡбрޑжሽǶᢀჸ ᏢಞԖځख़ाޑғӸሽॶǴΨ֎Ї΋٤ࣽᏢৎ׫Ε೭ঁሦୱޑࣴزǶّϞ੤ቺ܎

ȐAlbert Banduraȑӧ 1977 ԃ܌ගрޑȨޗ཮Ꮲಞ౛ፕȩȐSocial learning theoryȑǴ ගٮΑ΋ঁࣴزޑख़ाୖԵаϷ౛ፕ୷ᘵǶ

ӧ੤ቺ܎ޑޗ཮Ꮲಞ౛ፕύǴჹՉࣁޑಞள௦ڗ׳ቶݱ܄ޑᢀᗺȐBandura;

1977/1995ȑǴᙁൂٰᇥǴ੤ቺ܎ᇡࣁՉࣁޑಞளࡽڙᒪ໺Ӣનޑज़ڋǴΞڙࡕϺ

࿶ᡍᕉნޑቹៜǴࡐᜤஒٿޣϩ໒ǶΓޑፄᚇՉࣁЬाࢂ٩ᒘࡕϺಞளޑǴԶᇡ

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ޕתᄽΑ΋ঁࡐख़ाޑӢનǶՉࣁಞளԖٿᅿόӕޑၸำǴ΋ᅿࢂ೯ၸޔௗ࿶

ᡍǴᕇளՉࣁᆶϸᔈ่݀۶Ԝ໔ᜢ߯ޑၸำǴᆀࣁȨ೯ၸϸᔈޑ่݀܌຾ՉޑᏢ ಞȩǹќ΋ᅿࢂӧޗ཮௃ნΠǴᢀჸޣȐobserverȑ೯ၸȸᢀჸȹҢጄޣ

ȐdemonstratorȑޑՉࣁԶಞளՉࣁޑၸำǴᆀࣁȨ೯ၸҢጄޣ܌຾ՉޑᏢಞȩǶ

ಃ΋ᅿՉࣁᕇளБԄջࣁঁᡏᏢಞǴӧ೭ᅿБԄ΢Ǵϸᔈޑ่݀ϐ܌аჹᏢಞࢂ

ԖᔅշޑǴЬाࢂӢࣁѬගٮΑΟᅿЬाфૈǴϩձࢂૻ৲ǵ୏ᐒǵᆶቚமфૈǶ ӧૻ৲фૈ΢ǴঁᡏૈவՉࣁϸᔈᆶ၀Չࣁ܌஥ٰޑ่݀ԶᕇளૈࡰᏤΠ΋ԛՉ

୏ޑૻ৲ǹӧ୏ᐒфૈ΢ǴӃ߻ޑ࿶ᡍ٬ளঁᡏჹܭࢌ໨Չ୏ޑ่݀ૈԖႣޕޑ གྷႽǴԶ٬ளঁᡏ཮ѐᒧ᏷୺Չ܈ό୺Չ၀Չ୏ǹӧቚமфૈ΢ǴΑှՖᅿՉ୏

ૈ஥ٰՖᅿၿ፞Ǵჹܭ၀Չ୏ࢂցૈ೏ߦ຾܈׭ڋࢂख़ाޑǶಃΒᅿՉࣁᕇளБ ԄջࣁᢀჸᏢಞǴӧ೭ᅿБԄ΢Ǵૻ৲ǵ୏ᐒǵᆶቚம೭٤фૈ߾ૈӧঁᡏόሡ ᒃيѐ୺Չ၀Չ୏ջૈ೸ၸᢀჸձΓޑՉ୏൩ૈᕇளǴΨӢԜ೭ኬޑᏢಞБԄǴ ჹΓᜪεӭኧޑՉࣁᕇளࢂख़ाޑǶ੤ቺ܎ᇡࣁǴᢀჸᏢಞࣁ΋ೱុޑᐕำǴԖ ѤঁЬाᐕำǴϩձࣁݙཀᐕำǵߥ੮ᐕำǵՉࣁᇙ೷ᐕำǵϷ୏ᐒᐕำǶݙཀ ᐕำࢂᢀჸᏢಞวғޑಃ΋؁ǴӧݙཀᐕำύǴҢጄޣՉ୏ҁيޑ੝ቻ܈மࡋǵ ᢀჸޣၸѐޑ࿶ᡍǵаϷᢀჸޣکҢጄޣϐ໔ޑᜢ߯฻ፏӭӢન཮ቹៜ๱Ꮲಞޑ ਏ݀ǶӧᢀჸᏢಞޑߥ੮ᐕำǴᢀჸޣૈஒ܌ᢀჸډޑ٣ҹᙯϯࣁࢌᅿ಄ဦࠠ

ԄǴջ٬Ңጄޣόӆр౜ǴΨૈ೸ၸ၀಄ဦࣁ൞ϟԶஒ܌ᢀჸډޑૻ৲੮ӧอය

૶Ꮻ܈ߏය૶ᏫύǶӧՉࣁᇙ೷ᐕำǴᢀჸޣሡाע૶Ꮻύޑ಄ဦᙯඤԋ፾྽ޑ ՉࣁǴ೭΋ၸำੋϷډ΋سӈᇡޕޑکՉࣁޑᏹբǴКӵჹ΋ՉࣁӵՖ߄౜ޑᇡ ޕǹૈჸ᝺ԾيՉ୏ᆶ૶ᏫύҢጄޣޑՉ୏ϐ໔ޑৡ౦٠຾Չঅ҅Ƕӧ୏ᐒᐕำ

΢Ǵᢀჸޣჹܭ܌ᢀჸޑ٣ҹࢂցڀȨයࡑȩǴ܈ޣा؃ᢀჸޣѐ߄౜ޑ௃ნࢂ

ցӝ፾ܭᢀჸޣ౜ӧޑރᄊǴࣣቹៜ๱ᢀჸޣࢂցૈ୼࿶த߄౜р܌ಞளޑՉ ࣁǶ೭Ѥঁᐕำޑൻᕉ຾ՉǴ٬ளനಖՉࣁૈ٩ᒘ๱ᢀჸᏢಞޑБԄԶᕇளǶ

੤ቺ܎ 1977 ԃޑޗ཮Ꮲಞ౛ፕǴаȨᇡޕȩޑཷۺٰඔॊΓᜪޑՉࣁᕇளǴ ԶΓᜪޑε೽ϩՉࣁёаᙖҗᢀჸᏢಞ೭ኬޑБԄٰᕇளǶӵ݀ाჹΓᜪޑՉࣁ

(12)

ᕇளᆶځङࡕޑЈ౛ᐒڋԖ׳ӭޑΑှǴךॺ൩Ѹ໪ाჹᢀჸᏢಞ೭ኬޑᏢಞБ ԄԖ׳ӭޑࣴزᆶ௖૸Ƕҗܭတઓ࿶ࣽᏢӧ߈жޑזೲஙଆǴೱ஥Ӧߦԋޗ཮ઓ

࿶ࣽᏢޑᑫଆǴӢԜȨᢀჸᏢಞȩङࡕޑઓ࿶ᐒڋ೴ᅌԋࣁ߈жઓ࿶ࣽᏢৎॺո Κགྷѐᑍ௖ޑ዗ߐ᝼ᚒǴՠҗܭаΓᜪࣁڙ၂ޣததԖፏӭज़ڋǴा׳ుΕӦѐ

௖઩ϣӧޑЈ౛ᐒڋࣗԿܭઓ࿶ᐒڋ೿ࢂόܰޑǶ࣬ၨϐΠǴ୏ނኳࠠගٮΑߡ ճ܄ǵܰᏹբ܄ǵаϷӢ݀௢ፕ࣬ჹᙁൂ฻฻ӳೀǴӢԜӵ݀ૈ୼೸ၸ୏ނኳࠠ

ٰࣴزᢀჸᏢಞ೭ኬޑ᝼ᚒǴό໻Ԗշܭ҂ٰᔈҔډΓᜪޑᢀჸᏢಞǴΨԖշܭ ךॺѐΑှ୏ނޑᇡޕૈΚǶ

ಃΒ࿯ʳ ӣ៝౜ϞӸӧܭ୏ނي΢ޑᢀჸᏢಞࣴز

ၸѐଞჹόӕނᅿᆶόӕՉࣁ೿ԖፏӭࣴزҾკѐ௖૸ᢀჸᏢಞޑ᝼ᚒǴԶ ёૈቹៜᢀჸᏢಞޑӢનΨԖӣ៝Ўക၁ಒӦ૸ፕ (Zentall, 2006)Ƕ೸ၸӣ៝ᆶ

૸ፕ೭٤ࣴزǴךॺёаᏢಞډӵՖ೛ी΋ঁ࣬ჹֹ᏾ޑჴᡍѐࣴزวғܭ୏ނ ي΢ޑȸᢀჸȹᆶȸᏢಞȹǶ

൘ǵᆶғӸԖᜢޑᢀჸᏢಞՉࣁ

΋ǵᏢಞ၀Ӟϙሶ१ނ

ଞჹ೭ᜪޑࣴز᝼ᚒǴӭъࢂࣴز٩ᒘ༘᝺ޑ୏ނࣁЬǴനத٬Ҕޑ୏ނኳ

ࠠࢂȨαښୃӳޑޗ཮໺ሀբ཰ȩȐsocial transmission of flavor preferences paradigmȑǶ၀բ཰ޑ୷ҁჴᡍำׇࣁǺ૽ግ΋ঁҢጄޣѐ१ҔӃ߻வؒӞၸޑ ढ़ғ१ނࡕǴᡣҢጄޣᆶќ΋ᢀჸޣ࣬ೀ΋ࢤਔ໔ǴӆஒҢጄޣڗр٠ᡣᢀჸޣ ჹٿᅿढ़ғ१ނٰ຾Չᒧ᏷Ƕᢀჸޣӧٿᅿढ़ғ१ނޑᒧ᏷΢໼ӛᒧ᏷ҢጄޣӞ ၸޑ१ނǴ೭ኬޑ่݀᛾ܴᢀჸޣӧढ़ғޑރݩΠ໼ӛճҔΑ਒Γޑ࿶ᡍٰ຾Չ

(13)

१ނᒧ᏷ǶԖࣴزஒԜ཰Ҕٰࣴز୏ނᢀჸᏢಞϐߏය૶ᏫᐕำǴว౜εႵ ȐLong-Evans ratȑѝሡᆶҢጄޣอኩ 30 ϩដޑϕ୏ࡕ൩ૈ׎ԋჹҢጄޣමӞၸ ޑ१ނԖ܌ୃӳǴջ٬ҢጄޣςόӧǴ೭ኬޑᏢಞਏ݀ᗋࢂૈ࡭ុ΋ঁД (Galef

& Whiskin, 2003)ǶӕኬճҔ೭ኬޑբ཰ǴӧаৎႵȐMus domesticusȑࣁ୏ނኳ

ࠠޑࣴز߾ࡰрǴԜᅿᢀჸᏢಞ཮ڙډޗ཮่ᄬޑፄᚇࡋԶ೏ቹៜޑǶ྽Ңጄޣ

ࢂᢀჸޣޑηж܈ޣࢂᆶᢀჸޣԃស࣬ӕޑќ΋ሟ܄Ǵ೭ᅿ૶ᏫёаӧҢጄޣς όӸӧࡕ 4 ঁλਔᗋૈߥ੮ǹՠӧ 24 λਔࡕǴѝԖᆶᢀჸޣԃស࣬ӕޑሟ܄Ң ጄޣ܌೷ԋޑ૶Ꮻਏ݀ᗋӸӧǶӵ݀ஒҢጄޣޑኧໆቚӭǴ΢ॊٿᅿόӕҢጄޣ

܌೷ԋޑ૶Ꮻਏ݀ǴёаӧҢጄޣςόӸӧࡕ 4 ঁλਔᗋૈߥ੮ǹՠࣣܭ 24 λ ਔࡕ࣮όډ૶ᏫӸӧޑ߄౜ (Choleris, Guo, Liu, Mainardi, & Valsecchi, 1997)Ƕ ΒǵᏢಞব္ёаפډ१ނ

ଞჹ೭ᜪޑࣴز᝼ᚒǴӭъࢂࣴز٩ᒘຎ᝺ࣁЬޑ୏ނǶӧԜ᝼ᚒ΢٠ؒԖ

΋ঁ੝ۓޑࣴزኳԄǴՠ୷ҁޑᏹ׋БԄջࣁǺᡣᢀჸޣѐᢀჸҢጄޣӵՖפډ १ނǴฅࡕӆᡣᢀჸޣԾՉѐ൨פ१ނǶ೯தᢀჸޣ཮ᒧ᏷Ҕд܌ᢀჸၸޑБԄ

܈ᒧ᏷Ңጄޣ܌ᒧޑӦᗺٰפ൨१ނǶ (΋) аങᜪࣁኳԄ

೸ၸങᜪޑࣴزǴόѝว౜дॺёа೸ၸᢀჸձޑҢጄޣޑ߄౜ٰפډ१ނ ޑ܌ӧǴᗋว౜ፏӭቹៜᢀჸᏢಞޑӢનǶࣴزࡰрǴђКങȐPoecilia reticulataȑ

ୃӳၟᒿԖ࿶ᡍǵዕ஼ޑǵឦܭεიᡏޑҢጄޣ (Lachlan, Crooks, & Laland, 1998)ǶᗨฅᢀჸޣୃӳၟᒿԖ࿶ᡍޑҢጄޣǴՠό߄Ңૈ೷ԋؼӳޑᏢಞਏ݀Ƕ

ࣴزࡰрǴᢀჸޣၟᒿ߄౜ؼӳޑҢጄޣǴѝӧ΋໒ۈ߄౜΢Кၟᒿ߄౜ৡࠂޑ ҢጄޣٰளӳǹՠډΑࡕය໘ࢤǴϸԶ཮ᡂԋၟᒿ߄౜ৡޑҢጄޣޑᢀჸޣӧᏢ ಞೲࡋ΢Кၨז (Swaney, Kendal, Capon, Brown, & Laland, 2001)Ƕ཮р౜೭ኬޑ

౜ຝǴࣴزޣᇡࣁǴࡐёૈࢂӢࣁ߄౜ؼӳޑҢጄޣϐՉ୏Ǵߦ٬Αᢀჸޣஒݙ ཀΚܫӧቹៜ߄౜ޑᜢᗖՏ࿼܈ڈᐟނ΢ǴԶ٬ளᢀჸޣӧ߃ۈ໘ࢤૈ߄౜ၨ

ӳǹ೭ᅿቹៜݙཀΚޑਏ݀Ξᆀࣁ୔ୱ܄மϯȐlocal enhancementȑ܈ڈᐟނம

(14)

ϯȐstimulus enhancementȑǶ࣬ჹٰᇥǴၟᒿ߄౜ৡࠂҢጄޣǴᗨฅ΋໒ۈݙཀ ΚޑᘍܫؒԖКၨזܫډ҅ዴՏ࿼΢ǴՠҗܭόؼҢጄޣ୏բКၨᄌǴᡣᢀჸޣ

৒ܰၟᒿǴԶЪԖ׳ӭᐒ཮ёа୔ϩԖਏՉࣁᆶคਏՉࣁǴӢԜ೷ԋࡕය໘ࢤޑ ؼӳ߄౜ǹ೭ᅿਏ݀Ξᆀࣁё୔ϩ܄ޑၟᒿՉࣁȐdiscriminated following behaviorȑǶќѦǴᗨฅᢀჸޣ഻៿ၟᒿεიᡏǴԶ߄౜рӵӕΓᜪШࣚύதـޑ வ౲ȐconformityȑǴՠ྽ᢀჸޣคݤӧຎ᝺΢࣮ډԾρޑიᡏ܌ӧਔǴيೀܭε იᡏޑᢀჸޣᏢಞ߄౜߾Кيೀܭλიᡏޑᢀჸޣٰளৡǹᗨฅӆࡋܭຎ᝺΢ૈ

࣮ډԾρޑიᡏ܌ӧਔǴيೀܭεიᡏޑᢀჸޣϐ߄౜Ξ཮Кيೀܭλიᡏޑᢀ ჸޣٰளӳ (Day, MacDonald, Brown, Laland, & Reader, 2001)Ƕ೭ኬޑ౜ຝΨས ҢΑவ౲ᗨฅԖշܭ୏ނှ،ढ़ғୢᚒǴՠࡐёૈ཮ቹៜཥڻՉࣁޑౢғǶவ΢

ॊϐࣴز೿ᇥܴΑᢀჸᏢಞჹܭ୏ނӧפ൨१ނ΢ࢂख़ाЪڀቹៜ܄ޑǴόၸΨ ԖࣴزࡰрǴᢀჸޣ٠ߚ΋ښޑ٬Ҕ਒Γ࿶ᡍǶӵ݀ঁᡏޑ࿶ᡍ٠όሡाбрϼ εжሽਔǴКଆ٬Ҕ਒Γ࿶ᡍǴᗋࢂ཮٩ᒘঁᡏ࿶ᡍǹՠ྽٬Ҕঁᡏ࿶ᡍሡбр εжሽਔǴ߾཮ᒧ᏷٩ᒘ਒Γ࿶ᡍ (Kendal, Coolen, & Laland, 2004)Ƕӧΐڈങ ȐPungitius pungitiusȑޑࣴزΨว౜Ǵ྽ᢀჸޣᇡࣁ٬ҔԾي࿶ᡍǴԋфᕇள१ ނޑёૈ܄ၨଯਔǴКଆ٬Ҕ਒Γ࿶ᡍǴᗋࢂ཮٩ᒘঁᡏ࿶ᡍǹ྽ᢀჸޣᇡࣁ٬

ҔԾي࿶ᡍǴԋфᕇள१ނޑёૈ܄ၨեਔǴ߾཮٩ᒘ਒Γ࿶ᡍǶନԜϐѦǴ྽

ঁᡏޑ૶ᏫࢂຫΦϐ߻ޑ࿶ᡍǴΨ཮फ़եᢀჸޣ٬ҔԾي࿶ᡍޑёૈ܄ǴԶׯࣁ ٩ᒘ਒Γ࿶ᡍ (van Bergen, Coolen, & Laland, 2004)Ƕ

(Β) аܲᙝࣁኳԄ

೸ၸᢀჸᏢಞԶӧ೔१Չࣁ΢ள੻ǴၸѐᇡࣁᢀჸޣᆶҢጄޣࣁӕ΋ဂᡏǴ ཀջၨዕ஼ޑӕՔǴځਏ݀ၨӳǶόၸԖࣴزࡰрǴε໳ီȐBombus impatiens CRESSONȑӧᢀჸΑձޑணҤޑҢጄޣϐ߄౜ǴΨ཮଺рᆶҢጄޣ΋ኬޑᒧ᏷

(Worden & Papaj, 2005)Ƕ೭ኬޑ่݀ᇥܴΑǴόӕޑဂᡏΨૈ೸ၸᢀჸᏢಞٰᕇ ளӳೀǴӆԛᡍ᛾ᢀჸᏢಞڀഢΑᄽϯ΢ޑሽॶǶόၸ၀ࣴزΨᢀჸډǴ೭ኬޑ ਏ݀ջ٬ࢂҗΓπޑኳᔕҢጄޣΨૈၲډ࣬ӕޑਏ݀Ǵ՟ЯᇥܴΑ೭ኬޑՉࣁ߄

(15)

౜ǴёૈѝሡाൂપޑݙཀΚᐕำวғջёֹԋǶ

ມǵᆶ܄ᆶᕷ़ԖᜢޑᢀჸᏢಞՉࣁ

ଞჹ೭ᜪޑࣴز᝼ᚒǴӭъࢂࣴز٩ᒘຎ᝺ޑ୏ނࣁЬǶ೭٤ࣴز೯த٬Ҕ

؃ଽ܈Ҭଛϕ୏ኳԄǶ၀ኳԄޑ୷ҁБݤջࣁǺᡣᢀჸޣѐᢀჸ΋໢΋ሟวғ؃

ଽ܈Ҭଛϕ୏٣ҹǴϐࡕӆᡣᢀჸޣѐჹٿଫόӕܭᢀჸޣ܄ձޑڈᐟނ଺ᒧ

᏷Ǵځύ΋ଫؒԖ೏ᢀჸޣᢀჸၸԖวғၸ؃ଽ܈Ҭଛϕ୏٣ҹǴќ΋ଫ߾ࣁ೏

ᢀჸޣᢀჸၸԖวғ࣬ӕ٣ҹޑҢጄޣǶӧԜ᝼ᚒΠ܌วғޑᢀჸᏢಞǴςόӆ

ࢂᢀჸޣᢀჸൂ΋Ңጄޣ୺Չࢌ໨Չ୏ԶவύᏢಞډҢጄޣޑՉ୏ǴԶࢂٿঁҢ ጄޣޑՉ୏೏ᢀჸᆶᏢಞǴӢԜ೭ኬޑࣴزǴჹܭዴᇡᢀჸޣ܌߄౜рٰޑՉࣁ

ࢂցᏢಞаϷவব္Ꮲಞډ၀Չࣁࢂሡाߚதᙣ཈ޑǶ

΋ǵаങᜪࣁኳԄ

ӧђКങޑࣴزύࡰрǴሟ܄ᢀჸޣ཮ୃӳම೏ᢀჸၸԖᆶځѬሟ܄຾Չၸ

؃ଽϕ୏٣ҹޑ໢܄ങǴЪᢀჸޣࢂᇡள၀໢܄ω຾ՉԜᒧ᏷ޑ (Dugatkin, 1992)ǶќѦǴᢀჸᏢಞޑቹៜΚόѝૈቹៜሟ܄ჹ໢܄ޑᄊࡋǴΨૈׯᡂচҁ ჹ໢܄ޑᄊࡋǶࣴزว౜ԁ៧޸吃ങȐPoecilia latipinnaȑӧᢀჸډচҁ܌ୃӳ ޑ໢܄೏ځѬሟ܄ܔ㶥ǴΨ཮ӧϐࡕܔ㶥၀໢܄ǴԶ೭ኬޑᄊࡋׯᡂߚᒿᐒวғ ޑ (Witte & Ueding, 2003)Ƕ׳ᡋΓЪԖ፪ޑࢂǴᢀჸᏢಞ܌೷ԋޑਏ݀ջ٬ӧό ӕࠔᅿي΢Ψૈว౜ǶࣴزࡰрǴሟ܄ԁ៧޸吃ങ཮ୃӳӃ߻೏ᢀჸډԖჹќ΋

ᅿ௦ค܄ғ෗БԄЪࣁόӕࠔᅿޑሟ܄٥ଭᇽԁങȐPoecilia Formosaȑ຾Չ؃

ଽՉࣁޑ໢܄ԁ៧޸吃ങ (Schlupp, Marler, & Ryan, 1994)Ƕ ΒǵаചᜪࣁኳԄ

(΋) ሟ܄ᢀჸޣ

೸ၸВҁ᜺᜷ȐCoturnix japonicaȑ܌ளډ่݀ᆶӧങᜪي΢܌ளډޑࣴز

(16)

่݀࣬྽ᜪ՟ (Galef & White, 1998)ǴନΑሟ܄ᢀჸޣѝሡᢀჸډ໢܄ᆶќ΋ଫ ሟ܄ઠӧ΋ଆ൩ૈჹ၀໢܄ౢғୃӳ (White & Galef, 1999a)Ǵ೭ኬޑ่݀Ψᇥܴ

Αᢀჸډȸ੝ۓՉ୏ȹޑวғჹሟ܄ᢀჸޣനࡕޑᒧ᏷٠ߚࢂѸाޑచҹǶќ ѦǴሟ܄ჹܭ໢܄ޑୃӳᗨฅ཮٬ள೏ୃӳޣ٬ځڙѫޑᐒ౗ၨଯǴՠᢀჸᏢಞ ޑਏ݀ૈቚуό೏ୃӳޣ٬ሟ܄ڙѫޑᐒ౗ගଯ (Persaud & Galef, 2005)ǶКၨ

Ԗ፪ޑࢂǴӵ݀໢܄ي΢܌ڀഢޑ੝ቻᆶሟ܄ᢀჸޣӃ߻ᢀჸډќ΋ଫමᆶځѬ ሟ܄Ҭଛၸޑ໢܄ي΢ޑ੝ቻ࣬ӕǴ߾ሟ܄ᢀჸޣΨ཮ჹڀԖ࣬ӕ੝ቻޑ໢܄Ԗ

܌ୃӳǴջ٬Ӄ߻ؒـၸ၀໢܄ (White & Galef, 2000)Ƕ (Β) ໢܄ᢀჸޣ

ӧᢀჸޣࣁ໢܄ਔ߾ளډᆶ΢ॊሟ܄่݀࣬ϸޑჴᡍ่݀Ǵ໢܄ϸԶ௨Ѿම ᢀჸډ೏ձޑ໢܄؃ଽၸޑሟ܄Ƕ೭ኬޑਏ݀ሡԖᢀჸډ؃ଽՉࣁޑр౜ωӸ ӧǴΨᇥܴΑᢀჸډȸ੝ۓՉ୏ȹޑวғჹ໢܄ᢀჸޣനࡕޑᒧ᏷ࢂѸाޑచҹ (White & Galef, 1999b)Ƕ

ୖǵᆶٛᑇԖᜢޑᢀჸᏢಞՉࣁ

ଞჹ೭ᜪޑࣴز᝼ᚒǴЬाӧࣴزঁᡏӵՖ೸ၸᢀჸᏢಞٰಞளૈᗉխӒᓀ วғޑՉ୏Ƕ೭ᅿ᝼ᚒޑࣴزБݤؒԖ΋ঁڰۓኳԄǴ೯தࢂᡣᢀჸޣѐᢀჸҢ ጄޣӵՖ଒ᚆ΋ঁܴᡉڀ࠶ુ܄ޑڈᐟނ܈ᕉნǶᗨฅૈ଒ᚆ၀࠶ુޑБԄёૈ

ԖࡐӭᅿǴՠᙖҗҢጄޣޑҢጄǴᢀჸޣஒёזೲᏢಞډࢌᅿ଒ᚆБԄǶ

΋ǵаങᜪࣁኳԄ

ӧђКങޑࣴزύว౜Ǵ྽Ңጄޣӧ౜൑ਔǴᢀჸޣ཮٬ҔҗҢጄޣٗᜐᏢ ಞٰޑၡጕٰזೲ଒ᚆ࠶ુ܄ڈᐟǹՠ྽ҢጄޣόӧਔǴᢀჸޣ܌٬Ҕϐ଒ᚆၡ ጕࠅёૈό཮ࡪྣҢጄޣ྽߃Ңጄޑٗኬѐ຾ՉǴϸԶᒧ᏷Αќ΋ᅿૈ׳זೲ଒

ᚆ࠶ુ܄ڈᐟޑၡጕ (Brown & Laland, 2002)Ƕ೭ኬޑ่݀όѝᇥܴᢀჸޣᙖҗ

(17)

ᢀჸᏢಞளډӳೀǴΨӆԛསҢΑவ౲཮ቹៜཥڻՉࣁޑౢғǶ ΒǵаܲᙝࣁኳԄ

ӧᐋᖧȐNemobius sylvestrisȑޑࣴزύว౜ǴᐋᖧӧᢀჸԖ࿶ᡍޑӕᜪ߄౜

рӵՖၧᗉഁᇐ຅ȐPardosa spp.ȑޑՉࣁࡕǴᢀჸޣΨ཮ճҔӕኬБԄٰၧᗉ

ഁᇐ຅ޑр౜Ǵջ٬Ңጄޣςόӆр౜Ǵ೭ኬޑᏢಞ߄౜ᗋࢂёа࡭ុ 24 λਔ (Coolen, Dangles, & Casas, 2005)Ƕ೭ኬޑ่݀ᇥܴΑջ٬Ⴝᐋᖧ೭૓ၸ๱ߚဂۚ

ғࢲޑғނΨૈճҔᢀჸᏢಞٰᕇளӳೀǶ Οǵаথ٢ᜪࣁኳԄ

(΋) а㖺ᏁᜪࣁኳԄ

ࣴزࡰрǴλႵȐCF-1ȑӧᢀჸӕᜪ೏ڀװᔐ܄ޑ૨ηѕࠈਔ཮р౜ੵภό

௵གϯȐanalgesiaȑޑ౜ຝǹԶᢀჸޣӧ 24 λਔࡕၶډ၀੝ۓᜪࠠޑ૨ηਔǴ ջ٬၀૨ης೏ܘନ֎ᆅԶคݤ೷ԋ࠶ુǴΨ཮ӵӕҢጄޣ΋૓଺рஒيᡏশ຾

პ਑ޑၧᗉБԄǴՠᢀჸޣ܌߄౜рޑ೭ᅿՉࣁ٠ό཮ӧᢀჸᏢಞ໘ࢤύр౜ӧ ᢀჸޣي΢ǶନԜϐѦǴᗋ཮р౜ڋऊ܄ੵภό௵གϯȐconditioned analgesiaȑ ޑ౜ຝ (Kavaliers, Choleris, & Colwell, 2001)Ƕࣴزᗋว౜Ǵ೸ၸ೭ኬޑȸᢀ ჸȹǴ཮೷ԋᢀჸޣޑҜ፦✉Ȑcorticosteroneȑ΢ϲǹԶᢀჸޣӧ 24 λਔࡕၶ ډ၀੝ۓᜪࠠޑ૨ηਔǴջ٬၀૨ης೏ܘନ֎ᆅԶคݤ೷ԋ࠶ુǴΨ཮ౢғڋ ऊ܄Ҝ፦✉΢ϲޑ౜ຝ (Kavaliers, Colwell, & Choleris, 2003)Ƕ೭٤่݀೿ᇥܴ

ΑǴᢀჸޣޑዴӧჹวғӧӕᜪي΢ޑ٣ҹ຾ՉᢀჸǶ (Β) аᒗࣁኳԄ

ࣴزύा؃ᢀჸޣǴӧόޔௗаަᡏௗ᝻ډҢጄޣޑ௃׎ΠǴᢀჸҢጄޣޑ ߄౜ࡕૈᏢಞډ྽᠋ډᖂॣૻဦ൩ाၢډጃηޑќ΋ᜐٰၧᗉႝᔐޑр౜Ƕ่݀

ᡉҢǴᢀჸޣಞள଒ᗉՉࣁޑೲࡋזܭؒԖ຾ՉၸᢀჸᏢಞޑᒃي࿶ᐕޣǶԖ፪ ޑࢂǴᢀჸޣӧ຾ՉᢀჸᏢಞ૽ግ߻Ǵම࿶ӧႝᔐጃύ೏ႝᔐၸ΋ԛ (John, Chesler, Bartlett, & Victor, 1968)ǴӢԜࡐёૈӃ߻ޑॄӛ࿶ᡍǴ٬ளᢀჸޣჹܭ วғӧӕՔي΢ӕኬޑᎁၶ཮׳௵ག܈׳ݙཀǴԶ٬ளᢀჸᏢಞޑਏ݀೏уமǶ

(18)

(Ο) аФࣁኳԄ

ԖࣴزࡰрǴϏഝ಻ើФȐDanish Friesian cowȑ཮᎞߈ྕࢋޑྣ៝ޣǹჹ ಉኪޑྣ៝ޣ߾߄౜рᇻᚆޑՉࣁǶόၸᢀჸޣ೸ၸᢀჸӕՔ೏аόӕБԄჹࡑ ٠คݤӢԜԶᏢಞډόӕྣ៝ޣϐྣ៝БԄǴᗨฅӧय़ჹӕՔࢂ೏аྕࢋޑБԄ ჹࡑਔǴᒿ๱ӕՔᆶྣ៝ޣޑຯᚆຫ߈ǴΨ཮߄౜рຯᚆᅌ߈ޑՉࣁǶ೭ኬޑ่

݀ᗨฅؒԖࡰрФૈ຾ՉԜᅿࠠԄޑᢀჸᏢಞǴՠځՉࣁ཮ڙډӕՔޑՉࣁቹៜ (Munksgaard, DePassill, Rushen, Herskin, & Kristensen, 2001)Ƕ

(Ѥ) аญηࣁኳԄ

ӧԃᇸޑࡡݞญȐmacaca mulattaȑޑࣴزύว౜Ǵᢀჸޣ೸ၸӧቹТύᢀ ჸډӕᜪჹೂ߄౜р্܂ޑϸᔈࡕǴ߾চҁ߄౜рόឨ܂ೂޑԃᇸࡡݞญǴΨ཮

ӧϐࡕय़ჹೂޑр౜ਔ߄౜р্܂ޑϸᔈǹՠӧӕኬޑᏹ׋ΠǴѝࢂஒቹТύޑ

ೂඤԋ޸ԚǴࠅό཮೷ԋԃᇸࡡݞญѐჹ޸Ԛ଺р্܂ϸᔈ (Cook & Mineka, 1990)Ƕ೭ኬޑᏢಞᐒڋ܈೚ёаҔᢀჸڋऊȐobservational conditioningȑٰှញǴ ཀջᢀჸޣڙӕᜪޑ৮ឨ௃ᆣ܌གࢉǴԶΞௗ๱य़ჹډচҁຎࣁύ܄ڈᐟނޑ

ೂǴӢԶౢғΑᙁൂᖄ่ᏢಞǶόၸൂൂаᙁൂޑᖄ่ᏢಞٰှញёૈΞό୼ֹ

ӄǴӢࣁԃᇸࡡݞญ٠ό཮Ꮲಞډा্܂޸Ԛ೭ᜪค໾্܄ޑނࠔǶ

စǵኳү

೭ᜪ᝼ᚒޑࣴز཮ा؃୏ނѐ୺Չ΋سӈޑՉ୏ำׇǴӭъࢂࣴز٩ᒘຎ᝺

ޑ୏ނࣁЬǶࣴزҞޑࣁ೸ၸᢀჸᏢಞǴёа٬ளᢀჸޣӧ୺Չ΋ঁཥޑՉ୏ׇ

ӈਔ׳ࣁזೲ܈ҍၨϿޑᒱᇤǴᙖԜٰࣴزӸӧܭ୏ނي΢ޑኳүȐimitationȑǶ ೯தჴᡍύ๏ϒ୏ނԋф୺ՉՉ୏ޑዛ፞ࢂ१ނǴԶჴᡍޣ཮ᡣᢀჸޣೀܭ΋ঁ

ବᎧޑރᄊǴځҞޑӧቚуᢀჸޣޑᏢಞ୏ᐒᆶ߄౜୏ᐒǶZentall ӧ 2006 ԃޑ ӣ៝Ўകύᇡࣁा࠹ᆀ୏ނޑ߄౜ࣁኳүਔǴሡाૈ௓ڋȨޗ཮ߦ຾ȩȐsocial

(19)

facilitationȑǴཀջᢀჸޣᆶҢጄޣೀӧӕ΋ᕉნ܌Ꮴठޑࢲ୏Κ΢ϲǴԶࢲ୏Κ ޑ΢ϲёૈ཮೷ԋՉ୏೏ଽฅֹԋǶᗋाૈ௓ڋӃ߻܌ගၸޑȨ୔ୱ܄மϯȩǵȨڈ ᐟނமϯȩǵᆶȨᢀჸڋऊȩǴаϷȨᇨวᏢಞȩȐaffordance learningȑǴཀջᢀჸ ޣ೸ၸҢጄޣޑՉ୏ԶᏢಞډڈᐟނ܈ᕉნޑфૈǶനࡕǴाዴᇡ೭٤Չ୏ಔԋ ӧচҁ୏ނي΢ࢂόӸӧҭ܈ӧؒԖҢጄޣޑՉ୏ҢጄΠǴᢀჸޣԾฅр౜၀Չ

୏ޑёૈ܄ࢂࡐեޑȐZentall, 2006ȑǶ೭٤Ӣનޑ௓ڋǴ٠ߚࡰ੿҅ޑኳүό཮

Ԗ೭٤ӢનӸӧǴԶࢂࡰ࠹ᆀࢌՉ୏ࣁ੿҅ޑኳүਔǴѸ໪ा௨ନҔ೭٤Ӣનջ

ૈֹӄှញ၀Չ୏р౜ޑёૈǶӧࣴز୏ނޑኳүǴؼӳޑ௓ڋಔ೛ीБԄЬा

ёϩٿᅿǺ΋ᅿࢂᅰёૈஒ΢ॊ܌ගӢન௨ନ௞ǹќ΋ᅿࢂȨᚈӛޑ௓ڋำׇȩ Ȑbidirectional control procedureȑǴ၀բ཰ा؃୏ނӧࢌ΋ނᡏ΢୺Չٿᅿόӕϸ ᔈǴԶόӕಔޑᢀჸޣஒ࣮ډҢጄޣ୺ՉόӕޑϸᔈǴӵԜ٬ள۶ԜջࣁჹБޑ

௓ڋಔǶ

΋ǵஒёૈӢન௨ନ௞

(΋) аᒗࣁኳԄ

ӧᒗޑࣴزύว౜Ǵा؃ᢀჸޣӧόޔௗаަᡏௗ᝻ډҢጄޣޑ௃׎ΠǴᢀ ჸډҢጄޣޑ߄౜ࡕૈᏢಞډ྽᠋ډૻဦջа߻ဌඓѐᓸఎηٰᕇள१ނǶ೭ኬ ޑՉ୏ा؃ޑ੝ձϐೀӧܭ୏ނѝૈаȸ߻ဌඓȹѐᓸఎηǴӢԜࢂа੝ۓيᡏ

೽ՏֹٰԋՉ୏ޑǶࣁΑዴᇡᢀჸޣ੿ޑԖᏢಞډ၀Չ୏Ǵ٠Ъ೭ኬޑᏢಞࢂፄ ᚇޑᐒڋǴӢԜࣴزύ೛ीΑٿঁ௓ڋಔǴځ΋ࢂᢀჸޣᢀჸډҢጄޣѝࢂ᎞߈ ሺᏔ൩Ԗ१ނӞǴ೭ኬޑ೛ीࢂ௨ନݙཀΚᐕำวғ܌ᏤठՉࣁౢғޑёૈ܄Ǵ Ψ൩ࢂ௓ڋ୔ୱ܄܈ڈᐟނமϯǹځΒࢂᢀჸޣᢀჸډҢጄޣόᆅҔҺՖБԄᓸ ఎη೿Ԗ१ނӞǴ೭ኬޑ೛ीࢂӧዴᇡ၀Ꮲಞёૈόѝࢂᙁൂޑᖄ่ᏢಞǴКӵ ᢀჸڋऊǶ่݀ᡉҢǴԖᢀჸډҢጄޣҔ߻ဌඓѐᓸఎηޑ཮КќѦٿಔ௓ڋಔ ߄౜р׳ӭ҅ዴᆶ׳זೲޑϸᔈ (John et al., 1968)Ƕ

(20)

(Β) а㖺ᏁᜪࣁኳԄ

ӧλႵ

Ȑ

C57BL/6J

ȑ

ޑࣴزύǴा؃ᢀჸޣӧόޔௗаަᡏௗ᝻ډҢጄޣ ޑ௃׎ΠǴᢀჸҢጄޣޑ߄౜ࡕૈᏢಞډӃډሺᏔ਒ᜐᓸΠ΋ߎឦעЋ٬ᐒᜢ௴

୏Ǵ྽१ނӢԜԶ௞ပډሺᏔޑΠቫܜஔࡕǴाາډሺᏔޑ҅߻БஒܜஔѺ໒ٰ

ڗள१ނ೭΋سӈፄᚇޑՉ୏Ǵ೭ኬޑፄᚇՉ୏ࢂҗวғӧٿঁόӕՏ࿼΢ޑٿ

ঁόӕՉ୏ࡪྣ΋ۓׇӈᜢ߯܌ಔԋޑǶ၀ࣴزӧᢀჸᏢಞ໘ࢤᆶᢀჸޣޑՉࣁ

ෳ၂໔ӼකΑ΋ঁਔ໔ޑۯ᪓Ǵ೭ኬޑ೛ीࢂࣁΑ௓ڋޗ཮ߦ຾Ǵёа௨ନอኩ ࢲ୏Κ΢ϲԶ೷ԋՉࣁр౜ޑёૈ܄Ƕ่݀ᡉҢ྽ᢀჸޣԖᢀჸډҢጄޣޑՉ

୏ǴКଆคݤᢀჸډҢጄޣޑλႵǴૈ೯ၸ೭໨ፄᚇբ཰ޑКٯ΢ၨଯ (Carlier

& Jamon, 2006)Ƕ Βǵᚈӛޑ௓ڋำׇ

(΋) а㖺ᏁᜪࣁኳԄ

ӧаεႵ

Ȑ

hooded Lister rat

ȑ

ࣁኳԄޑࣴزύǴ཮૽ግҢጄޣஒ΋ਥఎη ۳ѓ௢܈۳Ѱ௢ǴԖ΋ъޑᢀჸޣஒ཮࣮ډҢጄޣעఎηӛѰ௢Ǵќ΋ъ߾࣮ډ ఎη೏ӛѓ௢Ƕ่݀ᡉҢᢀჸޣ཮߄౜рдॺ܌ᢀჸډޑՉ୏ǶࣁΑ௨ନԴႵࢂ

ճҔఎηර๱܊໔ύόӕय़ӛᕅᏛٰ౽୏଺ࣁጕ઩ǴӢԜࣴزޣӧᢀჸޣޑᢀჸ ໘ࢤ่״ࡕ཮ஒఎηޑᘍܫՏ࿼౽ډќ΋य़ᕅᏛǴ٬ளఎηޑ౽୏Бӛவচҁޑ Ѱѓᡂԋѝૈ߻ࡕǶ่݀ӧෳ၂໘ࢤਔᢀჸޣᗋࢂ߄౜р܌ᢀჸډޑՉ୏

(Heyes & Dawson, 1990)Ƕ (Β) аചᜪࣁኳԄ

а㖺ᏁᜪࣁኳԄ཮ౢғ΋٤ୢᚒǴջคݤ௨ନԴႵޑ߄౜ёૈѝࢂ΋ᅿȨᇨ วᏢಞȩ܈ޣԴႵёૈճҔූ੮ܭఎη΢ޑښၰ྽଺ጕ઩ǶࣁΑှ،ӵԜୢᚒǴ

ࣴزޣճҔӕኬޑБԄࡼෳӧ٩ᒘຎ᝺ޑ୏ނي΢ǴӵВҁ᜺᜷ǴӢԜ൩ό཮Ԗ ښၰූ੮ޑୢᚒǶᢀჸޣஒӧᢀჸᏢಞය໔ᢀჸډߐ೏ҢጄޣරόӕБӛ౽୏Ƕ ჴᡍύቚуќ΋௓ڋಔǴᢀჸޣӕኬᢀჸډߐරόӕБӛ౽୏ǴѝࢂԜ౽୏όࢂ

(21)

Ңጄޣ܌ᏤठޑǶ่݀ᡉҢǴᢀჸޣ཮٩܌ᢀჸډߐޑ౽୏БԄѐՉ୏ǹՠӧ௓

ڋಔ΢߾ό཮ǶӢԜȨᇨวᏢಞȩҭคݤֹӄှញᢀჸޣ܌߄౜рٰޑՉࣁ (Akins, Klein, & Zentall, 2002)Ƕ

ಃΟ࿯ʳ ޗ཮܄ૻ৲᠍ڗ

ኳүΨࢂᢀჸᏢಞޑ΋ᅿǴόၸኳүமፓޑࢂᢀჸޣ೸ၸᢀჸԶಞளҢጄޣ ޑȸՉ୏ȹǴՠᢀჸޣځჴёаѝࢂ೸ၸᢀჸҢጄޣޑՉ୏Զಞளȸૻ৲ȹǶၸ ѐჹ୏ނޑᢀჸᏢಞՉࣁࣴزǴӭࢂӧ௖૸୏ނᙖҗᢀჸᏢಞޑБԄٰᕇளࢌᅿ Չ୏Ǵՠӵ݀ाჹᢀჸᏢಞ೭ኬޑᐕำԖ׳ӭޑΑှǴךॺόૈ۹ౣќ΋ᅿಞள ޑϣ৒Ǻૻ৲Ƕाࣴز೭᝼ᚒǴ܈೚೸ၸࣴز୏ނԾฅวғޑޗ཮܄ᝡݾՉࣁǴ ёаᡣךॺჹ೭ᅿȸૻ৲ȹಞளޑᢀჸᏢಞԖ׳ӭޑΑှǶ

൘ǵวғܭޗ཮܄ᝡݾύޑޔௗຑ՗ᆶ໔ௗຑ՗

ӧғނ໔Ǵӕᜪ໔۶Ԝϕ୏ᝡݾࢂࡐี᚞ѳதޑǴځҞޑεӭࢂࣁΑݾၗ

ྍǴ೭٤ၗྍёૈࢂ१ނޑٰྍǵҬଛޑ៾Κǵ܈ޣ෈৲ӦޑጄൎǶՠӵࣁΑݾ

ၗྍǴ؂ԛ೿Їଆᐟਗ਼ޑѺରǴჹ୏ނٰᇥࢂڀ॥ᓀ܄Ϸ໾্܄ޑǴӢࣁᐟਗ਼ޑ ѺରёૈᏤठڙ໾ࣗԿܭࢂԝΫǶ܌а୏ނӧᝡݾύሡჹӕᜪޑᏯରૈΚ຾Չຑ

՗Ǵ྽Ѻରၸำύຑ՗рჹЋૈΚКԾρமၸ೚ӭǴ൩όा੮ᠰܭၗྍݾڗǴᔈ

྽ᅰזܫకᏯରќ൨рၡǶՠঃऩሡा೸ၸ؂ԛჴሞᏯର࿶ᐕωૈຑ՗ჹЋޑૈ

ΚǴԶคݤ૶ڗ೏਋௳ၸޑ௲૽Ǵ߾ڙ໾ޑ॥ᓀᗋࢂ཮ӸӧǶӢԜ୏ނाૈᏢಞ ډӧ੝ۓ࿶ᡍΠ຾ՉঁᡏӆᇡȐindividual recognitionȑǴཀջૈ୔ϩჹЋࢂցම

࿶਋௳ၸԾρǴӵ݀ӆᇡрჹЋම࿶਋௳ၸԾρǴ߾ᔈ྽ᒧ᏷όᏯԶଏǶၸѐӧ

௶ճ٥ঊႵȐMesocricetus auratusȑޑࣴزύ൩ࡰрǴ໢܄ঊႵӧ೏਋௳ࡕǴӵ

(22)

ӆࡋޔௗय़ჹ࣬ӕޑჹЋਔ཮όᏯԶ଒ǹନԜϐѦǴჹ၀੝ۓჹЋޑ૶Ꮻёа࡭

ុԿϿΎϺ (Lai & Johnston, 2002)Ƕՠ٣ჴ΢Ǵջ٬೭ኬૈᏢډ௲૽Ǵёૈᗋࢂ

ڀԖ΋ۓޑ॥ᓀ܄ǴӢࣁሡाᒃي࿶ᐕᏯରωૈֹԋຑ՗Ƕ٣ჴ΢ǴӵӕӃ߻ჹ วғܭ୏ނي΢ޑᢀჸᏢಞՉࣁ܌຾Չޑӣ៝ǴᢀჸᏢಞࢂڀԖᄽϯ΢Ϸ୏ނғ Ӹ΢ख़ाޑሽॶǴӢԜךॺёа௢ෳǴନΑ೸ၸޔௗޑᒃي࿶ᐕٰ຾ՉእЋຑ՗

ѦǴᔈ၀Ӹӧ๱ૈ೸ၸᢀჸдΓޑ࿶ᡍٰᕇளૻ৲Ǵ຾ԶၲډᕭอእЋຑ՗ᐕำ ޑБԄǶ٣ჴᡉҢǴ೸ၸഁӦᢀჸᆶჴᡍ࠻ࣴزǴ೭ᅿ೸ၸ਒Γޑ࿶ᡍٰၲډૻ

৲ᕇளޑБԄࢂӸӧޑǴᆀࣁȨޗ཮܄ૻ৲᠍ڗȩȐeavesdroppingȑǶ

ޗ཮܄ૻ৲᠍ڗ΋ຒ೏٬Ҕٰඔॊ୏ނ೸ၸᢀჸќѦٿଫӕᜪޑϕ୏Զவ ϕ୏ύௗԏډૻ৲ޑ΋ᅿБԄ (McGregor, 1993)ǶEavesdrop ೭ঁ୏ຒࢂҗ eavesdripᙯᡂԶٰǶEaves ࢂࡂᛞޑཀࡘǹԶ drip ࡰளࢂᅀΠǴӢԜ eavesdrip ӝଆٰࡰޑ൩ࢂࡂᛞஒߘНௗՐޑ೭ᅿБԄǴԶӵԜБԄ཮೷ԋߘН໩๱ࡂᛞޑ

ྎࢬډࡂѦѐǶҗܭђжהޣୂ᠋ਔ཮ஒԾρॹ௠ӧࡂᛞǴ൩ӳႽߘНࢬΠޑኬ ηǴࡕٰ೭ᅿୂ᠋ޑՉࣁΨ൩೏ᆀࣁ EavesdroqǴԶ೭ኬޑཷۺ൩೏ۯ՜ډ೭ᅿ

ࣴزૻ৲ᕇளޑ᝼ᚒύǶޗ཮܄ૻ৲᠍ڗޑ၁ಒۓကࣁǴ೸ၸᢀჸќѦٿଫӕᜪ ޑϕ୏Ǵ຾Զவϕ୏ޑૻဦύѐ๧ڗрૻ৲٠ӧ҂ٰѐ٬Ҕϐ (McGregor, 1993;

McGregor & Peake, 2000)Ƕӧϕ୏ύ܌х֖ޑૻ৲ȐinformationȑǴКБӵᏯରૈ

ΚǴࠔ፦Ǵ୏ᐒ฻฻Ǵ೭٤ૻ৲όࢂҗૻဦȐsignalȑҁي܌ಔԋǴԶࢂӸӧܭ ೭٤ૻဦ࡛ሶ೏٬Ҕӧ೭ϕ୏ϐύǶ೯தाബ೷΋ঁޗ཮܄ૻ৲᠍ڗޑᕉნǴന ᙁൂޑБԄ൩ࢂᡣӕ܄ձ܈ޣόӕ܄ձޑᢀჸޣѐᢀჸќѦٿଫӕ܄ձޑ୏ނ

຾Չᝡݾ܄ϕ୏٣ҹǶाዴᇡᢀჸޣࢂցςᕇளૻ৲Ǵ߾ሡाѐෳໆᢀჸޣӧᢀ ჸ٣ҹวғϐࡕǴय़ჹ܌ᢀჸډޑٿঁҢጄޣޑϸᔈࢂցԖ܌όӕǴӵ݀Ԗόӕ ջё࠹ᆀᢀჸޣςճҔޗ཮܄ૻ৲᠍ڗޑБԄᕇளૻ৲ǶӧԜ᝼ᚒ΢а୏ނࣁኳ

ࠠޑࣴزςԖऩυЎ᝘่݀ǴԖ٤ࣴز಄ӝޗ཮܄ૻ৲᠍ڗޑۓကǹՠԖ٤߾ό

಄ӝǴՠ೸ၸΑှ೭٤ࣴزύޑࣴز฼ౣᆶ܌ளډޑ่݀ǴஒԖշܭךॺჹ೭΋

᝼ᚒޑ௖૸Ƕ

(23)

ມǵޗ཮܄ᝡݾύଞჹޗ཮܄ૻ৲᠍ڗӧ୏ނي΢ޑࣴز

΋ǵഁӦࣴز

ԐයഁӦࣴزࡰрǴ໢܄आવ໵㚁ȐAgelaius phoeniceusȑӧي਒ޑᎃۚ೏

ѦእߟΕԾৎሦӦਔǴ྽ߟΕޣࢂ΋ଫόڀװᔐ܄ޑܿऍࢪЦ៙㒣ȐTyrannus

tyrannusȑǴᢀჸޣߟҍ၀ᎃۚሦӦޑՉࣁ൩൳ЯؒวғǹՠӵߟΕޣࢂќ΋ଫڀ

װᔐ܄ޑ໢܄आવ໵㚁Ǵ྽ᎃۚӧय़ჹ೭ᅿѦእߟΕਔ܌๏ϒޑװᔐၨ১ਔǴ߾

ᢀჸޣߟҍԜ੝ۓᎃۚሦӦޑՉࣁ൩཮ၨதวғ (Freeman, 1987)Ƕ೭ኬޑ่݀

՟Я໔ௗᇥܴ๱໢܄आવ໵㚁ёૈ೸ၸȸᢀჸȹٰᏢಞᎃۚޑ០እૈΚǶ Βǵჴᡍ࠻ࣴز

(΋) аങᜪࣁኳԄ 1. ໢܄ᢀჸޣ

ӧа໢܄ᐁᛥରങȐBetta splendensȑࣁኳԄޑࣴزύǴࣴزޣࢂᡣᢀჸޣ ѐᢀჸٿଫ໢܄ങӧϕ୏୔ϣ຾Չ 5 ϩដޑᝡݾ܄ϕ୏Ƕ೸ၸ೸ܴ႖݈ஒ۶Ԝ୔

႖໒ٰǴ٬ளᢀჸޣᆶٿଫҢጄޣӅΟଫ໢܄ങǴคݤӧަᡏ΢຾Չޔௗௗ᝻Ǵ ՠૈ೸ၸຎ᝺ᆶ༘᝺ޑБԄٰ຾Չૻဦᆶૻ৲ޑ໺ሀǶӧ೭ኬޑჴᡍᏹ׋ΠǴࣴ

زޣԖᢀჸډǴ྽ᝡݾ܄ϕ୏຾ՉϐਔǴᢀჸޣ཮य़ჹԖ٣ҹวғޑٗঁБӛǴ ӢԜࣴزޣᇡࣁᢀჸޣԖӧჹ٣ҹ຾ՉᢀჸǶӧᢀჸᏢಞ໘ࢤ่״ࡕǴࣴزޣ཮

ஒϕ୏ύޑᒡৎȐloserȑڗрǴ੮ΠគৎȐwinnerȑᆶᢀჸޣǴ႖Ϻ߾຾ՉՉࣁ

ෳ၂໘ࢤǶՉࣁෳ၂ᕉნࢂќ΋ঁཥᕉნǴគৎᆶᢀჸޣ཮೏ᒿᐒϩԋٿಔǴ΋

ಔᢀჸޣ཮ᆶচٰ܌ᢀჸၸޑគৎҢጄޣଛჹǹќ΋ಔᢀჸޣ߾཮ᆶќ΋ଫؒـ ၸՠᡏख़ᆶځ࣬ৡόӭޑគৎҢጄޣଛჹǶ؂ঁଛჹ཮೏ܫӧϕ୏୔ϣǴӧؒԖ

೸ܴ႖݈ޑ௃׎Π຾Չޔௗ܄ޑᝡݾ܄ϕ୏Ǵࣴزޣ߾ஒϕ୏ރݩ૶ᒵΠٰǶࣴ

ز่݀ࡰрǴᢀჸޣόፕനࡕࢂയࢂॄǴӧ،ۓाᐟᏯ܈ޣाܫక΢Ǵӧᆶዕ஼

ޑគৎҢጄޣ຾Չϕ୏Кଆᆶढ़ғޑគৎҢጄޣ຾Չϕ୏ޑ೭ٿᅿ௃ნΠǴᢀჸ

(24)

ޣӧᆶ߻ޣޑϕ୏΢ϐՉ୏،฼཮׳זೲǴᡉҢӃ߻ޑᢀჸૈ෧ϿእЋૈΚຑ՗

ޑਔ໔Ǵٰᡣঁᡏ׳זೲ଺рղᘐ (Johnsson & Ækerman, 1998)Ƕ೭ኬޑࣴز่

݀КଆഁӦᢀჸளٰޑ่݀Ǵ׳Ԗᐒ཮᛾ܴᢀჸޣԖӧჹᝡݾ܄ϕ୏٣ҹ຾Չᢀ ჸǶόၸǴջ٬ӵԜǴ၀ࣴزᗋࢂӢࣁჴᡍ೛ी΢ޑόֹഢԶคݤܴዴ᛾ܴᢀჸ ޣԖӧჹᝡݾ܄ϕ୏٣ҹ຾ՉᢀჸǴӢࣁڙ၂ޣරӛԖ٣ҹวғޑٗঁБӛǴё

ૈѝࢂӢࣁٗঁБӛԖӕᜪޑӸӧǶӕԃޑќ΋ঁࣴزύǴ߾଺Αၨ٫ޑ௓ڋٰ

ዴᇡԜ٣ǶࣴزޣஒჴᡍሺᏔϩԋᢀჸ୔ᆶϕ୏୔Ǵϕ୏୔ࢂӧᢀჸ୔ޑѰѓٿ ᜐǶᢀჸޣ཮೏Ӽ௨ᢀჸځύ΋ᜐޑϕ୏୔ύԖٿଫҢጄޣ೸ၸ೸ܴ႖݈ӧ຾Չ ໔ௗޑᝡݾ܄ϕ୏٣ҹǹќ΋ᜐ߾ᢀჸډٿଫҢጄޣӢࣁύ໔႖๱ό೸ܴ႖݈Զ

ؒԖӧ຾Չϕ୏ǶҗܭҢጄޣᆶᢀჸޣ܌ೀޑ୔ୱԖܴߝࡋ΢ޑৡ౦ǴӢԜᢀჸ ޣૈᢀჸډҢጄޣޑࢲ୏ǴՠԾيό೏Ңጄޣ๏ᢀჸډǶӧ೭ኬޑ೛ीΠǴ၀ࣴ

زࡰрǴ࣬ჹܭؒԖᝡݾ܄ϕ୏٣ҹวғޑ΋ᜐǴᢀჸޣჹܭԖวғᝡݾ܄ϕ୏

٣ҹޑٗ΋ᜐ཮ݙຎၨΦǴᡉҢΑᢀჸޣޑዴԖӧݙཀᝡݾ܄ϕ୏٣ҹޑวғǶ

ௗ๱ࣴزޣགྷዴᇡᢀჸޣࢂցவᝡݾ܄ϕ୏٣ҹޑᢀჸύಞளૻ৲ǴӢԜ௦ҔΑ ќ΋ᅿ೛ीǶᢀჸޣ཮೏Ӽ௨ᢀჸځύ΋ᜐޑϕ୏୔ԖٿଫҢጄޣ೸ၸ೸ܴ႖݈

ӧ຾Չ໔ௗޑᝡݾ܄ϕ୏٣ҹǹќ΋ᜐ߾Ӣό೸ܴ႖݈ߔ႖๱ǴᏤठคݤᢀჸډ ќѦٿଫҢጄޣӕኬӧ຾Չ໔ௗޑᝡݾ܄ϕ୏٣ҹǶᢀჸᏢಞ໘ࢤ่״ࡕ཮ଭ΢

຾ՉՉࣁෳ၂໘ࢤǴ܌ԖޑҢጄޣ཮ϩձ೏းӧ΋ঁ೸ܴ౯ηύǴа΋ԛ΋ଫޑ БԄ࿼Εᢀჸޣ܌ࡑޑ୔ୱϣǴࣴزޣ߾ෳໆᢀჸޣჹߟΕޣޑϸᔈǶӧ೭ኬޑ

೛ीΠǴ၀ࣴزࡰрǴ࣬ჹܭय़ჹ๱ᢀჸډᒡၸдΓޑΕߟޣǴᢀჸޣჹܭᢀჸ ډគၸдΓޑΕߟޣ཮߄౜р጗ᄌޑᖿ߈Ǵ٠Ъၨᄌωჹځ຾Չ࠶ᓵǹՠ೭ኬޑ ৡ౦܄ӧόᡣᢀჸޣԖᐒ཮ᢀჸډҢጄޣॺ܌຾Չޑᝡݾ܄ϕ୏٣ҹਔ൩ό཮

р౜Ƕ೭٤่݀ᡉҢΑڙ၂ޣޑዴԖӧჹӕᜪޑᝡݾ܄ϕ୏٣ҹ຾ՉᢀჸǴ٠வ ύಞளૻ৲ǴЪஒ٤ૻ৲଺ࣁࡕុᏯର൑ӝޑ߃ۈՉ୏٩Ᏽ (Oliveira, McGregor,

& Latruffe, 1998)Ƕჹܭዴᇡᢀჸޣ܌ಞளૻ৲ࢂٰԾܭӕᜪޑᝡݾ܄ϕ୏ϐ ύǴନΑճҔ୏ނ܌วғޑ੿ჴᏯରǴᗋԖࣴز٬ҔΑќ΋ᅿѯ֮ޑ೛ीٰዴᇡ

(25)

Ԝ٣ǶࣴزύǴࣴزޣᡣᢀჸޣൂБय़ᢀჸډϕ୏୔ԖٿଫҢጄޣǴA ᆶ B ೸ ၸ೸ܴ႖݈ӧ຾Չ໔ௗޑᝡݾ܄ϕ୏٣ҹǶѯ֮ޑࢂǴᝡݾ܄ϕ୏٣ҹύځჴࢂ

ٿٿ໢܄ങӧ۶Ԝ຾Չϕ୏ǴA ᆶ B аϷ C ᆶ DǶࣴزޣஒ B аϷ C ᗦᙒՐǴ

٬ளவᢀჸޣޑفࡋٰ࣮ǴӢࣁѝ࣮ډ A ᆶ DǴԶᇤаࣁࢂ A ᆶ D ӧ຾Չϕ୏Ƕ ӧᢀჸᏢಞ໘ࢤ่״ࡕǴࣴزޣ཮ϩձᡣᢀჸޣᆶ A ܈ B ຾Չ໔ௗޑᝡݾ܄ϕ

୏ǴӕኬෳໆᢀჸޣჹჹЋޑϸᔈǶӧ೭ኬޑ೛ीΠǴࣴزࡰрǴ࣬ჹܭय़ჹᢀ ჸډȸᒡၸдΓȹޑჹЋǴᢀჸޣჹܭᢀჸډȸគၸдΓȹޑჹЋ཮ၨזޑᖿ߈ аϷၨזޑჹځ຾Չ࠶ᓵǶҗܭӧ೭ኬޑ೛ीΠǴ٬ள੿ჴШࣚ܌วғޑ٣ҹᆶ ᢀჸޣ܌ޕ᝺ډޑ٣ҹࢂόӕޑǴӢԜᢀჸޣჹόӕჹЋ߄౜рόӕՉࣁǴѝૈ

ᘜӢܭ೭ٿঁჹЋӧᢀჸޣޑ౳Πޑ࣬ჹ߄౜܌Ꮴठޑ (McGregor, Peake, &

Lampe, 2001)Ƕ

2. ሟ܄ᢀჸޣ

ޗ཮܄ૻ৲᠍ڗΨૈၮҔӧሟ܄ϐଛଽᒧ᏷΢Ƕࣴزว౜Ǵሟ܄ᐁᛥରങӧ य़ჹԖวғᝡݾ܄܈ؒวғᝡݾ܄ϕ୏٣ҹ΢཮߄౜рόӕՉࣁǴԶჹܭԖวғ ᝡݾ܄ϕ୏٣ҹޑٿঁ໢܄ڈᐟނǴᢀჸޣ཮ᒧ᏷Ӄ᎞߈គৎҢጄޣǴ٠޸ၨӭ ਔ໔ݙຎаϷ߄౜рၨӭޑғ෗ՉࣁǴ՟ЯКଆᒡৎҢጄޣǴሟ܄ᢀჸޣКၨୃ

ӳគৎҢጄޣ (Doutrelant & McGregor, 2000)Ƕ (Β) аചᜪࣁኳԄ

1. ໢܄ᢀჸޣ

ࣁΑؼӳ௓ڋϕ୏ၸำаϷஒૻ৲ޑӸӧᆒዴ௓ڋӧϕ୏ύǴԖ΋ᅿჴᡍЋ ݤ൩ࢂճҔኞܫᏔѐኳᔕٿଫ໢ചᙖҗሳћٰ຾Չᝡݾ܄ϕ୏٣ҹǴԶ٣ҹ຾Չ ύǴ཮Ԗќ΋ଫ੿ჴޑ໢܄ചӧ਒຾ՉᢀჸǶࣴزࡰр໢܄εξഒȐParus majorȑ όፕࢂჹᝡݾ܄ϕ୏٣ҹวғޑᢀჸ྽Π܈ޣय़ჹаኞܫᏔኳᔕԋޑΕߟޣޑ ߟౣǴ೿཮߄౜рזೲᖿ߈ޑՉࣁǴԶЪࡕޣК߻ޣᗋٰள׳זǶჹܭΕߟޣޑ ߟౣǴ྽ԜߟΕޣӃ߻೏ᢀჸޣ᠋ډӧᝡݾ܄ϕ୏٣ҹύࢂឦܭۚܭ১༈ޑ΋

БǴ߾཮ჹѬԖၨϿޑӣᔈǴᡉҢᢀჸޣૈᙖҗᢀჸӕᜪ໔ޑ۶ԜሳћރᄊǴٰ

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ಞளӕᜪ໔ޑՏ໘ᜢ߯ (Peake, Terry, McGregor, & Dabelsteen, 2001)Ƕ 2. ሟ܄ᢀჸޣ

ӧаሟ܄ᐁᛥରങࣁኳԄ܌ளډޑ่݀ǴӧаВҁ᜺᜷ࣁኳԄ΢ࠅؒԖளډ

࣬ӕ่݀ǶࣴزࡰрǴӧᢀჸၸᝡݾ܄٣ҹޑวғࡕǴሟ܄ᢀჸޣ཮ᒧ᏷ۚΠ॥

ޑ໢܄ڈᐟނǶࣴزޣჹܭ೭่݀ޑှញࢂǴሟ܄ёૈाᗉխ೏ᏱԖଯװᔐ܄ޑ

໢܄ӧ຾ՉҬଛਔװᔐǹ၀ࣴزΨว౜Ǵຫڀᓬ༈ޑ໢܄Ǵຫ཮ӧ຾ՉҬଛਔ଺

р໾্܄ޑ୏բ (Ophir & Galef, 2003)Ƕ

ୖǵঁᡏ࿶ᡍჹޗ཮ᝡݾύޑޗ཮܄ૻ৲᠍ڗޑቹៜ

ᢀჸޣӧ຾ՉእЋຑ՗ϐਔǴၸѐ܌ᏱԖޑ਒Γ࿶ᡍ՟Я՞Ԗख़ाޑቹៜ ΚǶόၸࣴزΨว౜ǴᢀჸޣόѝёаճҔ਒Γ࿶ᡍǴᗋёаӝٳ٬Ҕঁᡏ࿶ᡍ ᆶ਒Γ࿶ᡍ೭ٿᅿόӕٰྍޑૻ৲Ƕ၀ࣴزӕኬճҔኞܫᏔѐኳᔕٿଫ໢܄εξ ഒᙖҗሳћٰ຾Չᝡݾ܄ϕ୏٣ҹǶჴᡍϩΟঁ໘ࢤǴӧಃ΋໘ࢤǴᢀჸޣȐSȑ

཮ௗڙঁᡏ࿶ᡍޑᏹ׋Ǵёૈࢂ೏਋௳ȐA > Sȑ܈ޣࢂᏯയߟΕޣȐA < Sȑǹӧ

ಃΒ໘ࢤǴᢀჸޣ཮ௗڙ਒Γ࿶ᡍޑᏹ׋ǴӢԜ཮ᢀჸډӃ߻ޑߟΕޣᆶќ΋ჹ Ћ຾Չᝡݾ܄ϕ୏ȐA > B ܈ A < BȑǹӧಃΟ໘ࢤǴӧಃΒ໘ࢤ܌ᢀჸډޑќ΋

ჹЋȐBȑஒ཮ߟΕᢀჸޣޑሦӦǶ่݀ࡰрǴᢀჸޣ೸ၸٿᅿόӕૻ৲ٰྍޑ ӝٳԶஒߟΕޣղᘐࣁόڀ࠶ુਔȐS > A > BȑǴ཮ჹ၀ߟΕޣԖၨϿޑӣᔈǹ ՠӵղᘐࣁڀ࠶ુ܄ȐB > A > SȑǴ܈ޣၗૻόىคݤΠղᘐਔȐS > A < B ܈ S <

A > BȑǴ߾཮Ԗၨӭޑӣᔈ (Peake, Terry, McGregor, & Dabelsteen, 2002)Ƕ೭ኬ ޑ౜ຝࢂ࣬྽Ԗ፪ޑǴӢࣁ೭ኬޑ߄౜຾΋؁སҢ๱ޗ཮܄ૻ৲᠍ڗ೭໨ૈΚว චډཱུठਔёૈ཮р౜Ȩሀ౽௢౛ȩȐtransitive inferenceȑǴӕਔᡉҢ೭ᅿ೏ຎࣁ ଯ฻ᇡޕޑ߄౜Ψૈ೸ၸᢀჸޣᏢಞޑБԄٰ຾ՉǶӧሦβങȐAstatotilapia burtoniȑޑࣴزύ൩ว౜Ǵᢀჸޣӧೱុᢀჸ΋ჹΞ΋ჹޑଛჹᝡݾ܄ϕ୏٣ҹ

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ࡕǴёૈૈ୼௢౛рჹЋ AǵBǵCǵDǵаϷ E ϐ໔Ӹӧ๱ A > B > C > D > E ޑՏ໘ᜢ߯ǶӢࣁջ٬ᢀჸޣؒᢀჸၸჹЋ A ᓬܭ E ܈ B ᓬܭ D ޑଛჹಔӝǴ Ψ཮߄౜рᇻᚆჹЋ A ՠᖿ߈ E ޑՉࣁ܈ޣᇻᚆ B ՠᖿ߈ D ޑՉࣁ (Grosenick, Clement, & Fernald, 2007)Ƕ

ಃѤ࿯ʳ ҁፕЎޑࣴز᝼ᚒᆶ฼ౣ

೸ၸޗ཮܄ૻ৲᠍ڗ೭ᅿᢀჸᏢಞޑБԄٰಞளૻ৲Ǵ٬ளঁᡏӧ٬Ҕ਒Γ

࿶ᡍਔǴӭΑ΋ᅿঁᡏԾՉᒧ᏷ޑޜ໔Ǵঁᡏёаᒧ᏷ा٬Ҕ܈ޣό٬ҔǴࣗԿ ӝٳځѬٰྍޑૻ৲ࡕӆ଺،ۓǴӢԜךॺᇡࣁ೭ኬޑᏢಞБԄࢂ΋ᅿଯ฻ᇡޕ

ૈΚޑ߄౜ǶќѦǴޗ཮܄ᝡݾӧғނ໔ࢂࡐதـޑϕ୏٣ҹǴࣴزวғӧޗ཮

܄ᝡݾϕ୏ύޑޗ཮܄ૻ৲᠍ڗࢂుڀғᄊਏࡋޑǶၸѐଞჹޗ཮܄ૻ৲᠍ڗ೭ ᅿࠠԄޑᢀჸᏢಞǴӭъࢂ٬Ҕങᜪ܈ࢂചᜪ଺ࣁ୏ނኳԄǴ௦ҔΓᜪаѦޑځ Ѭথ٢ᜪϐࣴز߾ࢂϿϐΞϿǶӢԜӵ݀ૈճҔ㖺Ꮑᜪ೭ኬ࣬ჹКၨଯ฻Ъௗ߈ ΓᜪޑނᅿǴࣗԿࢂ㖺Ꮑᜪჴᡍ୏ނǴٰࡌҥрཥޑޗ཮܄ૻ৲᠍ڗϐ୏ނኳ ԄǴନΑቚ຾ךॺჹόӕނᅿӧԜૈΚ΢ޑΑှѦǴΨԖշܭךॺճҔԜჴᡍ୏

ނኳԄѐ຾΋؁௖૸ᆶᙶమԜՉࣁङࡕޑတ೽ઓ࿶ᐒڋǶ

൘ǵࣴز᝼ᚒ

΋ǵޗ཮܄ૻ৲᠍ڗᆶ௶ճ٥ᅿঊႵ

௶ճ٥ঊႵ೭ᅿ㖺Ꮑᜪ୏ނǴ೏ᇡࣁԋԃࡕӧഁӦύࢂՉᐱۚޑғࢲΔӧჴ ᡍ࠻ύԋԃࡕΨӭаᐱҥޜ໔ޑБԄஒ۶Ԝϩ໒ႩᎦǶԋԃঊႵჹܭځѬΕߟ ޣڀԖଯװᔐ܄Ǵ཮ࣁΑٛፁԾρޑࢰҤᆶሦӦԶᆶځѬӕᜪวғᏯରǶόၸ ঊႵ٠ߚ΋рғ൩ࢂᐱۚǴ೯தӧᚆணࡕ܈ࢂࢌঁ໘ࢤࡕǴω໒ۈځᐱۚғࢲǴ

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ՠջ٬ՉᐱۚғࢲǴځࢲ୏ሦୱΨᆶځдӕᜪԖ΋ۓำࡋޑख़᠄ᆶϕ୏ǶԖࣴ

ز൩ࡰрǴᐱۚ܄ၨ১ޑԋԃ೑ࣔࠠঊႵȐPhodopus campbelliȑӧαښୃӳޑ ޗ཮໺ሀբ཰΢ޑᢀჸᏢಞਏ݀ၨӳǴՠ࣬ჹٰᖱᐱۚ܄ၨமޑԋԃ௶ճ٥ঊ Ⴕ߾ѝԖ྽Ңጄޣࢂᢀჸޣޑ҆ᒃǴ٠Ъᢀჸޣԃइऊ 4 ঁࢃයεਔωૈว౜

ԖᢀჸᏢಞޑਏ݀ǶόၸКଆόࢂவҢጄޣ቏္ڗрޑ१ނǴԋԃ௶ճ٥ঊႵ

ୃӳᒧ᏷வҢጄޣ቏္ڗрޑ१ނ (Lupfer, Frieman, & Coonfield, 2003)Ƕᇥܴ

ΑԃѴޑ௶ճ٥ঊႵڀԖ΋ۓޑᢀჸᏢಞૈΚǹԶԋԃޑঊႵǴӧᒧ᏷१ނ΢

ᗋࢂёа೸ၸᢀჸᏢಞٰಞளǴѝࢂӢࣁલЮᒃஏޗ཮ϕ୏ޑ୏ᐒ܈ࢂࣁΑᗉ խόѸाޑѺରǴӢԶफ़եΑ೭ᜪᢀჸᏢಞวғޑᐒ཮Ǵՠϝฅڀഢ΋ۓำࡋ ޑᢀჸᏢಞૈΚǶҗܭԋԃঊႵڀԖሦୱ܄ՉࣁǴӢԜ྽ၶډΕߟϐঊႵǴ൩ ёૈ཮วғѺରՉࣁǶ೭ᅿ੝܄ς೏ԋфճҔٰᡍ᛾р໢܄ঊႵࢂڀԖঁᡏӆ ᇡޑૈΚǴΨ൩ࢂૈӆᇡрѺគၸԾρޑ੝ۓእΓ (Lai & Johnston, 2002)Ƕӧ ଛଽᒧ᏷ϐᢀჸᏢಞࣴز΢ΨࡰрᢀჸᏢಞੋϷΑঁᡏӆᇡޑᐕำ (Dugatkin, 1992)ǶӢԜճҔԋԃ໢܄ঊႵԖሦӦٛᑇՉࣁǵёаӧჴᡍ࠻ൂᐱႩᎦᗉխό Ѹाυᘋǵу΢ڀԖૈঁᡏӆᇡޑૈΚǵаϷڀԖ΋ۓำࡋޑޗ཮ϕ୏ᆶᢀჸ ᏢಞૈΚ฻ᓬᗺǴԖёૈԖᐒ཮ёаӧԋԃঊႵ΢ᢀჸډޗ཮܄ᝡݾϕ୏ύޑ ޗ཮܄ૻ৲᠍ڗϐᏢಞБԄǴ٠຾΋؁ࡌҥჴᡍᏹբϐՉࣁኳࠠǶ

Βǵঁᡏ࿶ᡍᆶޗ཮܄ૻ৲᠍ڗ

ӧᒗޑᢀჸᏢಞࣴزύǴࣴزޣ೸ၸ٣Ӄ๏ϒᢀჸޣ΋ԛ೏ႝᔐ࿶ᡍࡕω຾

Չ଒ᗉႝᔐޑᢀჸᏢಞ૽ግǴԖว౜ᢀჸᏢಞਏ݀ޑр౜ (John et al., 1968)Ǵ՟

ЯӃ߻ޑॄӛ࿶ᡍ཮ߦ຾ࡕុޑᏢಞਏ݀ǶόၸǴ၀ࣴز٠ؒԖᏹ׋คӃ߻ঁᡏ

࿶ᡍΠޑᢀჸᏢಞ௃ნǴӢԜόዴۓӃ߻ޑॄӛ࿶ᡍࢂց཮ቹៜࡕុޑᢀჸᏢಞ ᐕำǴՠ೸ၸа໢܄εξഒࣁኳࠠӧޗ཮܄ૻ৲᠍ڗ᝼ᚒ΢܌଺ޑࣴزǴ໔ௗ࣮

ډᢀჸޣӃ߻ޑόӕঁᡏ࿶ᡍǴ཮ቹៜࡕុځ٬Ҕ਒Γ࿶ᡍޑБԄǴᗨฅ၀ࣴز ჹܭᢀჸޣӧ਒Γૻ৲ᕇளϐਔԖՖϸᔈ٠คගϷ (Peake et al., 2002)ǶӢԜ௢

ፕǴӃ߻ޑঁᡏॄӛ࿶ᡍǴٯӵޗҬ਋௳Ǵᔈ၀Ψ཮ჹวғܭޗ཮܄ᝡݾϕ୏ύ

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ޑޗ཮܄ૻ৲᠍ڗϐᢀჸᏢಞᐕำ೷ԋቹៜǴࣗԿቹៜډࡕុޑᏢಞ߄౜Ƕ ΟǵᓸΚ಻ᅟᆾǵᓸΚೀ౛ᆶޗ཮܄ૻ৲᠍ڗ

຾Չޗ཮܄ᝡݾϕ୏Ǵёૈ཮೷ԋ୏ނڙ໾ǴӢԜ೭ኬޑ٣ҹჹܭ୏ނٰᇥ ёૈ൩ࢂڀᓸΚޑǴԶيᡏӵՖჹ೭ኬޑᓸΚ຾Չ΋ঁӝ፾ޑೀ౛Ǵ൩ᡂԋࡐख़

ाЪࡐჴҔޑ᝼ᚒǶΠຎЫ-တࠟᡏ-๝Ҝ፦ືጕȐhypothamamic-pituitary-adrenal cortex axisȑǴӧথ٢ᜪ୏ނय़ჹᓸΚ௃ნ܈ڈᐟޑೀ౛ᆶ፾ᔈ΢תᄽ๱ख़ाف ՅǶϩݜᐟ๝΢ဏញܫનȐcorticotrophin-releasing hormone, CRHȑޑઓ࿶ϡՏܭ ΠຎЫ਒࠻ਡȐparaventricular nucleusȑޑ߻೽ǹ྽Ѧࣚޑڈᐟࢂڀ࠶ુ܄ޑǴ

ૻဦ཮࿶җག᝺Ꮤ۔໺ډΠຎЫǴڈᐟځϩݜᐟ๝΢ဏញܫનǹ၀ញܫન཮೸ၸ တࠟဏߐેس಍ډၲတࠟဏ߻ယǴ٬ځϩݜᐟ๝΢ဏҜ፦નȐadrenocorticotropic hormone, ACTHȑǴ೭٤๝΢ဏҜ፦ન཮຾΋؁ӆѐڈᐟ๝΢ဏҜ፦ӝԋ๝΢ဏ ᗐҜ፦ᐟનȐglucocorticoidȑ (Engelmann, Landgraf, & Wotjak, 2004)Ƕ೭٤ᐟન

೏ញܫډՈనύ໺ၲډӄيǴჹ୏ނӧय़ჹᆙ࡚٣ҹਔࢂख़ाޑǶ྽୏ނࡋၸᆙ

࡚٣ҹࡕǴॄӣ㎸ፓ௓ᐒڋ཮௴୏٠׭ڋΠຎЫ-တࠟᡏ-๝Ҝ፦ືጕޑࢲ୏Ǵٰ

ᗉխՈనύޑ๝΢ဏᗐҜ፦ᐟન֖ໆၸଯ (Sapolsky, Romero, & Munck, 2000)Ƕ ӢԜǴໆෳҜ፦✉Ȑcorticosteroneȑ܈Ҝ፦ᎇȐcortisolȑ೭٤๝΢ဏᗐҜ፦ᐟન ޑᡂϯ೏྽଺ෳໆ୏ނޑᓸΚೀ౛ϐख़ाғ౛ࡰ኱ (M÷stl & Palme, 2002)ǶԖࣴ

زࡰрǴ೏૨ηװᔐޑҢጄޣǴځՈనύޑҜ፦✉֖ໆ཮΢ϲǹԶӧ਒ᢀჸҢጄ ޣᎁၶװᔐޑᢀჸޣǴՈనύҜ፦✉֖ໆΨ཮΢ϲ (Kavaliers et al., 2003)ǴᡉҢ Αय़ჹڀ࠶ુޑڈᐟނόѝࢂ೷ԋঁᡏౢғᓸΚϸᔈǴᢀჸӕᜪᎁၶڀ࠶ુ܄ޑ ٣ҹΨ཮٬ᢀჸޣౢғᓸΚϸᔈǶԖࣴزҭࡰрǴ໢܄ঊႵӧᆶځѬӕᜪ຾Չ΋

ԛᝡݾ܄ϕ୏Π೏਋௳ǴՈనϣޑҜ፦ᎇ֖ໆ཮΢ϲ (Huhman, Moore, Ferris, Mougey, & Meyerhoff, 1991)ǶՠӧڀԖ೏਋௳࿶ᡍΠѐᢀჸӕᜪ຾Չӕኬ܄፦ޑ ϕ୏٣ҹǴࢂց೷ԋᢀჸޣޑҜ፦ᎇ֖ໆӢԜڙډቹៜǴ߾ࢂ΋ঁ҂ޕኧǴᗨฅ ԖࣴزࡰрವΟКլαჯങ ( Oreochromis mossambicus ) ӧؒԖҺՖ࿶ᡍΠѐ ӧᢀჸӕՔޑᝡݾ܄ϕ୏٣ҹਔǴᡏϣޑ 11-keto-࿐ڰ✉Ȑ11-keto-testosteroneȑ

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ک࿐ڰ✉ޑ֖ໆ཮ϲଯǴᡉҢᢀჸޣჹᝡݾ܄ϕ୏٣ҹ຾ՉᢀჸǴғ౛΢ࢂ཮ڙ ډቹៜޑ (Oliveira, Lopes, Carneiro, & Canario, 2001)Ƕ

๝΢ဏᗐҜ፦ᐟનޑኬҁڗளٰྍЬाёаϩԋߟΕ܄ᆶߚߟΕ܄ޑБ ԄǶӧߟΕ܄БԄ΢ǴЬाࢂ೸ၸ׀ᓉેǵ౳ᆂǵЈ᠌ऀڈ܈ޣᘐᓐٰ௦ளՈన ฅࡕ຾Չϩ݋ǹӧߚߟΕ܄БԄ΢Ǵ߾ࢂԏ໣୏ނޑᕨߡٰෳໆᕨߡϣϐ๝΢ဏ ᗐҜ፦ᐟનжᖴނ֖ໆǶ࣬ၨϐΠǴߚߟΕ܄БԄКၨ໔ௗЪኬҁᇆ໣઻Κ຤

ਔǴࢂҞ߻КၨόදၹՠΞКၨ፾྽Ҕܭ຾ՉᓸΚ࣬ᜢࣴزޑБԄǴচӢࢂߟΕ

܄௦ՈБԄԖځᝄख़ޑલᗺᆶज़ڋǴٯӵǺӧ຾Չ௦Ոਔሡज़ڋ୏ނޑՉ୏܈ޣ

຾ՉഞᎈǴՠӵԜ୏բёૈ཮٬ளՈనύ๝΢ဏᗐҜ፦ᐟન֖ໆቚуԶᏤठჴᡍ ਏ݀೏ԡࢉ (Pace & Spencer, 2005; Vahl et al., 2005)Ǵӵ݀ࣴزޣௗΠٰᗋགྷाᢀ ჸ୏ނӧځѬՉࣁբ཰΢ޑ߄౜Ǵ຾Չ௦Ոਔ܌೷ԋޑᓸΚёૈ൩཮ቹៜ୏ނޑ Չࣁ߄౜ǹќѦǴࣴزޣӵ݀གྷाᢀჸڙ၂ޣՈనύ๝΢ဏᗐҜ፦ᐟન֖ໆޑೱ

ុਔ໔ᗺᡂϯǴനதـޑբݤࢂ٬Ҕόӕಔޑڙ၂ޣӧόӕਔ໔ᗺѐ຾Չ௦ՈǴ ՠ೭ኬޑբݤ཮ሡा٬Ҕεໆڙ၂ޣЪคݤஒ಻ᅟᆾᡂϯ΢ޑঁᡏৡ౦܄๏Ե ቾ຾ѐǶќ΋Бय़Ǵ٬ҔߚߟΕ܄БԄᗨёှ،΢ॊޑલᗺϷज़ڋǴՠሡԵቾӵ Ֆᒧ᏷ᕨߡԏ໣ޑਔ໔ᗺǴӢࣁჴሞ΢ჴᡍᏹ׋ϐਔ໔ᗺᆶ಻ᅟᆾжᖴނӧᕨߡ ϣ೏ୀෳډϐਔ໔ᗺϐ໔཮Ԗ΋ࢤۯ᪓ਔ໔Ǵ܌аӵՖᒧ᏷ᕨߡԏ໣ޑਔ໔ᗺǴ

཮،ۓࣴزޣࢂցૈୀෳډჴᡍᏹ׋܌೷ԋޑᓸΚϸᔈϐਏ݀Ƕࣴزว౜Ǵӧ୏

ނي΢ݙ৔ 3H-Ҝ፦✉Ȑ3H-corticosteroneȑǴ྽ݙ৔ਔ໔ࢂวғӧλႵȐC57BL/6Jȑ ޑསຼයаϷߝຼයϐਔǴ܌ෳໆډޑۯ᪓ਔ໔ϩձࢂ 4 λਔаϷ 10 λਔѰѓ (Touma, Sachser, Möstl, & Palme, 2003)ǹՠӧεႵȐSprague-Dawley ratȑي΢߾

ࣁ 17 λਔѰѓ (Bamberg, Palme, & Meingassner, 2001)ǴᡉҢΑۯ᪓ਔ໔ޑᡂ౦

܄ࢂࡐεޑǴ཮ڙჴᡍࢂวғӧ΋ϺύޑবঁຼයύаϷ܌٬Ҕޑ୏ނࠔسԶԖ

܌ቹៜǶନԜϐѦǴ၀ࠔᅿޑᓸΚϸᔈૈց೸ၸ೭ᅿߚߟΕ܄מೌٰୀෳрǴΨ

ࢂ٬ҔԜБԄਔሡाԵቾޑ (Rupert, 2005)Ƕᆕӝа΢܌ॊǴךॺᇡࣁǴҗܭࣴ

زޗ཮܄ૻ৲᠍ڗޑ᝼ᚒǴሡाᡣ໢܄ঊႵௗڙೱុޑᢀჸᏢಞ૽ግаϷᏢಞਏ

(31)

݀ෳ၂ǴӢԜ೭ኬޑᐕำёаႣය཮࡭ុ΋ࢤόอޑਔ໔ǴӢԜӧόቹៜ୏ނܭ ೭ࢤਔ໔ϣޑᏢಞᐕำаϷ୏ނޑВதࢲ୏ΠǴ௦ҔߚߟΕ܄Бݤѐ௖૸ޗ཮܄

ૻ৲᠍ڗᐕำޑ຾Չύ܌ౢғޑᓸΚϸᔈёૈࢂၨӝ፾ޑǶ

ມǵਡЈୢᚒᆶှ،฼ౣ

ҁፕЎύԖΟঁЬाޑਡЈୢᚒǴਥᏵ೭٤ୢᚒǴךॺ೛ीΑѤঁჴᡍǺ

΋ǵࡌҥคঁᡏޗҬ࿶ᡍచҹΠޑޗ཮܄ૻ৲᠍ڗϐ௶ճ٥໢܄ঊႵኳࠠ

೸ၸჹޗ཮܄ૻ৲᠍ڗ᝼ᚒޑӣ៝ǴךॺΑှډǴᢀჸޣ཮ჹӕᜪ܌วғޑ ᝡݾ܄ϕ୏٣ҹ຾ՉȸᢀჸȹǴԶᢀჸޣ཮வ٣ҹύಞளȸૻ৲ȹǴ٠ஒૻ৲٬

ҔӧࡕុᏯର൑ӝϐύǴ଺ࣁ୔ϩڀόӕᏯରૈΚޑჹЋϐ٩ᏵǶӢԜǴӧךॺ

܌ࡌҥޑ௶ճ٥໢܄ঊႵኳࠠύǴךॺሡा᛾ܴᢀჸޣԖӧჹڬᎁޑӕᜪϕ୏٣ ҹ຾ՉᢀჸǴ٠வύಞளૻ৲Ƕ܌аӧჴᡍ΋ǴךॺᡣؒԖҺՖѺର࿶ᡍޑ໢܄

ڙ၂ޣѐᢀჸǺќѦٿଫ໢܄Ңጄޣ຾ՉѺରǹ܈ޣ႖๱៓ᆛ຾Չύ܄ޗҬϕ

୏Ǵ೭ٿᅿόӕ܄፦ޑ٣ҹǶฅࡕܭќ΋ঁ൑ୱύज़ڋҢጄޣޑՉ୏Ǵа΋ԛග ٮ΋ঁ੝ۓҢጄޣښၰޑБԄǴෳໆڙ၂ޣჹ੝ۓښၰޑϸᔈǶӵԜՉࣁෳໆБ ԄǴ٬ளךॺૈ୼ෳໆډڙ၂ޣ٩ၸѐჹ੝ۓښၰޑ૶Ꮻ܌߄౜рޑՉࣁϸᔈǶ ӧ೭ኬޑ೛ीϐΠǴךॺଷ೛ڙ၂ޣӵ݀ԖჹҢጄޣ܌଺ޑՉ୏຾ՉᢀჸǴ߾ό ӕ܄፦ޑϕ୏٣ҹ܈ޣόӕ܄፦٣ҹύޑҢጄޣǴ཮٬ளڙ၂ޣӧᢀჸ྽Πޑ܈

ޣࡕុ༘ᆪډ੝ۓҢጄޣښၰϐਔޑՉࣁ߄౜཮Ԗ܌όӕǶќѦǴଷ೛ڙ၂ޣԖ வᝡݾ܄ϕ୏٣ҹύಞளૻ৲Ǵ߾ڙ၂ޣ༘ᆪډϕ୏٣ҹύޑគৎҢጄޣϐښၰ

܌߄౜рޑՉࣁ཮όӕܭ༘ᆪډϕ୏٣ҹύޑᒡৎҢጄޣϐښၰ܌߄౜рޑՉ ࣁǶ

ΒǵঁᡏޗҬ਋௳࿶ᡍࡕޑޗ཮܄ૻ৲᠍ڗ

೸ၸჹᢀჸᏢಞᚒ᝼ᚒޑӣ៝ǴךॺΑှډǴঁᡏӃ߻ޑॄӛ࿶ᡍёૈ཮ቹ

(32)

ៜᢀჸޑᏢಞᐕำаϷࡕុޑᏢಞ߄౜ǴӢԜӧჴᡍΒǴךॺ཮ӧჴᡍ΋܌ࡌҥ рޑޗ཮܄ૻ৲᠍ڗϐჴᡍᏹբՉࣁኳࠠύуΕঁᡏޑޗҬ਋௳࿶ᡍǴᡣڙ၂ޣ ӧڀԖ΋ԛ೏਋௳࿶ᡍΠѐௗڙӕኬޑᢀჸᏢಞ૽ግаϷࡕុޑՉࣁෳ၂Б ԄǶ਋௳ڙ၂ޣޑჹЋ཮ᆶڙ၂ޣஒाᢀჸډޑҢጄޣࢂόӕޑڈᐟނǴ೭ኬޑ

೛ीБԄஒ٬ளڙ၂ޣӧࡕុޑᏢಞ߄౜΢Ǵځϸᔈଞჹޑஒ཮ࢂӃ߻ᢀჸډޑ ϕ୏٣ҹύޑ੝ۓҢጄޣǴԶߚଞჹӃ߻๏ϒ਋௳࿶ᡍޑჹЋǶ

Οǵޗ཮܄ૻ৲᠍ڗᐕำΠᆶࡕុޑૻ৲٬ҔਔޑᓸΚϸᔈ

ӧჴᡍΟύǴЬाࢂໆෳ໢܄ঊႵӧҁፕЎޑჴᡍᏹ׋ΠϐᓸΚϸᔈǴёϩ ԋჴᡍΟ A ᆶჴᡍΟ BǶࣁΑໆෳ೭ኬޑᓸΚϸᔈǴךॺςߚߟΕ܄БԄǴٰ

ୀෳڙ၂ޣޑᕨߡϣҜ፦ᎇжᖴނȐfecal cortisol metabolites, FCMȑ֖ໆǶךॺ ςΑှډӧ٬ҔߚߟΕ܄БݤਔǴሡԵቾ܌ᒧ᏷ޑᕨߡԏ໣ਔ໔ᗺǴӢࣁѬஒ،

ۓࢂցૈໆෳډ୏ނӢჴᡍᏹ׋܌ౢғޑᓸΚϸᔈǴҗܭၸѐӕኬаߚߟΕБݤ ӧ௶ճ٥ঊႵي΢௖૸ᓸΚ࣬ᜢ᝼ᚒޑࣴزѝԖ΋ጇǴ၀ࣴز௦ڗԏ໣٣ҹ߻ࡕ Ӛ 24 λਔϣڙ၂ޣ܌௨ܫрٰޑᕨߡ (Chelini, Souza, Cortopassi, Felippe, &

Oliveira, 2006)ǴځѬޑࣴز߾ӭаλႵ܈εႵ଺ࣁڙ၂ޣ (Bamberg et al., 2001;

Touma et al., 2003)ǶҗܭঊႵޑᡏࠠϟΕλႵᆶεႵϐ໔Ǵՠ߈՟ܭλႵǴࣁΑ ዴߥ܌ᒧ᏷ਔ໔ᗺૈୀෳډᓸΚϸᔈǴךॺ཮٩ჴᡍҞޑሡ؃Ǵ௦ڗ؂ 3 ঁλਔ ൩ԏ໣΋ԛ໢܄ঊႵ܌௨ܫрٰޑᕨߡ܈ޣ؂ 18 λਔϣ܌௨ܫрٰޑᕨߡǶࣁ ΑዴᇡҁፕЎύ܌٬ҔޑߚߟΕ܄ᕨߡϣҜ፦ᎇжᖴނୀෳݤૈໆෳډڙ၂ޣ ޑᓸΚϸᔈǴךॺӧჴᡍΟ AǴ٬ҔΑԜמೌѐໆෳ໢܄ঊႵӧڀԖόӕ܄፦ޑ

ঁᡏޗҬ࿶ᡍΠޑᓸΚϸᔈǴӧ೭ኬޑޗҬ࿶ᡍΠǴڙ၂ޣ཮ڀԖ΋ԛ೏਋௳ޑ

࿶ᡍ܈ޣ΋ԛᏯയќ΋ଫ໢܄ঊႵޑ࿶ᡍǶךॺ௢ෳǴӵ݀೭ኬޑמೌૈᔈҔӧ

໢܄ঊႵي΢٠ໆෳрځ܌ౢғޑᓸΚϸᔈǴ߾ၸѐ೸ၸߟΕ܄௦ՈБݤӧ໢܄

ঊႵي΢аᜪ՟ޑᏹ׋܌ளډޑ่݀ (Huhman et al., 1991)ǴΨૈӧჴᡍΟ A ύ аߚߟΕמೌٰளډᡍ᛾ǶӧჴᡍΒޑ೛ीΠǴךॺᡣڙ၂ޣᆶќ΋ଫ໢܄ঊႵ

຾Չᝡݾ܄ϕ୏ԶᏱԖ೏਋௳ޑ࿶ᡍǴӆᡣڙ၂ޣа਒ᢀޣޑيϩѐय़ჹឦܭӕ

(33)

ኬ܄፦ޑᝡݾ܄ϕ୏٣ҹޑวғǴךॺ௢ෳǴڙ၂ޣӧ೭ኬޑᏢಞᐕำύаϷࡕ

ុ༘ᆪډϕ୏٣ҹύޑគৎҢጄޣϐښၰਔǴࡐёૈࢂೀܭԖᓸΚޑރᄊ΢ǶӢ ԜӧჴᡍΟ BǴךॺ཮ճҔԜמೌѐୀෳჴᡍΒύޑڙ၂ޣϐᓸΚϸᔈǶ

(34)

ಃΒകʳ ჴᡍ׷਑ᆶБݤ

ಃ΋࿯ʳ ჴᡍ΋

ҁჴᡍޑҞޑࢂճҔঊႵڀԖϐሦୱװᔐՉࣁࣁኳԄǴᡣؒԖҺՖޗҬ࿶ᡍ ޑԋԃ໢܄ঊႵѐᢀჸќѦٿଫ໢܄ঊႵޑޗҬϕ୏ՉࣁǴᙖԜٰࣴزޗ཮܄ૻ

৲᠍ڗޑ౜ຝࢂցӸӧܭঊႵي΢ǴΨ൩ࢂ҂ԖޗҬ࿶ᡍޑᢀჸޣૈցӧᢀჸٿ ଫҢጄޣޑѺର٣ҹࡕǴჹ೭ٿଫҢጄޣ߄౜рόӕޑՉࣁǴᙖа௢ፕޗ཮܄ૻ

৲᠍ڗӵԜޑᢀჸᏢಞБԄǴӧԋԃঊႵي΢ޑዴӸӧǶନԜϐѦǴ೸ၸғ౛ࡰ

኱ٰᡍ᛾ᢀჸၸำύ܌೷ԋޑቹៜǶ

΋ǵڙ၂ޣ

ҁჴᡍ௦Ҕ໢܄௶ճ٥ঊႵȐMesocricetus auratusȑǴрғ 8 ຼࡕԾ୯ৎჴ ᡍ୏ނύЈᖼளǴаঁձႩᎦ᠈Ȑᆫ✊༟ጤ׷፦Ǵߏ 33ǵቨ 21ǵଯ 15 Ϧϩȑࣁ ႖ᚆൂՏޑ IVC س಍Ȑindividual ventilated caging systemȑႩᎦܭ୯ҥѠ᡼εᏢ Ј౛س୏ނ܊ϣǴ᠈ϣᎎ೛ҏԯఒპ਑Ǵ٠кϩٮᔈႩ਑ᆶ໯НǶჴᡍύ܌٬Ҕ ڙ၂ޣӅ 34 ଫǴႵសऊࣁ 16 ډ 20 ຼǶ୏ނ܊ϣ೛ۓࣁ 12Ǻ12 ॹᙯϐВڹຼයǴ སຼයவԐ΢ 8 ᗺ໒ۈǴྕࡋᆢ࡭ 22 ± 2ʚǴྒྷࡋᆢ࡭ӧ 50 %ѰѓǶ

ΒǵՉࣁໆෳᏔ׷

ჴᡍύ܌٬ҔޑՉࣁໆෳᏔ׷ӅԖ 2 ᅿǴϩձࣁᢀჸᏢಞጃȐobservational learning boxȑᆶ U ׎଎ࢭȐU mazeȑǶ

(΋) ᢀჸᏢಞጃ

٬ҔᢀჸᏢಞጃޑҞޑࢂբࣁᢀჸᏢಞ໘ࢤ૽ግϐҔǴᡣڙ၂ޣёаᢀჸќ Ѧٿଫ໢܄ঊႵޑϕ୏Ƕӵკ΋ A ܌ҢǴᢀჸᏢಞጃȐߏ 44 Ϧϩǵቨ 44 Ϧϩǵ ଯ 15 ϦϩȑҗᓸլΚ׷፦܌ᇙԋǴ΢БԖ೸ܴᓸլΚ݈ᙟᇂǴБߡჴᡍޣޑᢀ ჸǴ٠ᗉխ୏ނޑ଒ಥǹځϣճҔᓸլΚ݈ᆶқՅλ៓᠈Ȑߏ 8.5 Ϧϩǵቨ 8.5

(35)

Ϧϩǵଯ 13.5 Ϧϩȑஒ୏ނ۶Ԝ୔႖໒ٰǶᢀჸᏢಞጃёаϩࣁȨᢀჸ୔ȩǵȨϕ

୏୔ȩᆶȨ฻ࡑ୔ȩΟঁ೽ϩǶᢀჸ୔ࢂҗٿঁқՅλ៓᠈η٠௨܌ಔԋǴٿঁ

᠈ηύ໔ақՅό೸ܴ႖݈୔႖໒ٰǴ٬ளٿଫڙ၂ޣ໔۶Ԝคݤ࣮ډჹБǴ؂

ԛჴᡍԿӭٿଫڙ၂ޣёаϩձܫ࿼Ελ៓᠈ύӕਔ຾ՉᢀჸᏢಞǶϕ୏୔ࣁ΋

ߏБࠠ୔Ȑߏ 22 Ϧϩǵቨ 15 Ϧϩǵଯ 15 ϦϩȑǴёᡣٿଫ໢܄ҢጄޣӧԜ୔ϣ

຾Չϕ୏Ƕ฻ࡑ୔ࣁ΋Οفࠠޜ໔Ȑ਒ᜐ 15 Ϧϩǵۭᜐ 8.5 Ϧϩǵ௹ᜐ 17 Ϧϩǵ ଯ 15 ϦϩȑǴࢂᢀჸᏢಞ૽ግᗋ҂໒ۈ߻ǴҢጄޣޑኩࡑϐೀǶᢀჸᏢಞጃϣӚ

୔ޑ࣬ჹՏ࿼Ǻᆶᢀჸ୔࣬ᎃޑࢂϕ୏୔Ǵٿঁ୔໔аλ៓᠈ηҁيޑ៓ํᆛஒ ٿ୔႖໒ǹӧϕ୏୔ٿ਒ޑࢂ฻ࡑ୔Ǵ୏ނࢂҗٿ਒ޑ฻ࡑ୔຾Εϕ୏୔Ǵ฻ࡑ

୔ᆶϕ୏୔໔߾аё౽୏ԄқՅό೸ܴᓸլΚ݈႖໒Ƕϕ୏୔ޑѦൎᕅᏛ΢ᆶ᎞

ᢀჸ୔ޑѦൎᕅᏛ΢ӕኬڀϾࢰȐϾ৩ελࣁ 0.7 ϦϩȑǴаճ਻ࢬޑࢬ೯Ƕӧ ᢀჸ୔ࡕБޑᕅѦǴ߾းԖႝ୏ܜ॥৻Ǵܭჴᡍ຾Չύ໒௴Ǵ਻ࢬஒҗϕ୏୔ޑ ѦൎᕅᏛ຾Εϕ୏୔നࡕډၲᢀჸ୔ǹ೭ኬޑ೛ीஒ٬ளښၰޑ໺ሀࢂൂӛޑǴ ନΑ٬ڙ၂ޣ৒ܰᆪډҢጄޣޑښၰѦǴᗋૈᡣҢጄޣόܰᆪډڙ၂ޣޑښၰԶ

ૈ஑ݙܭ౜ӧ҅຾ՉޑޗҬϕ୏΢ǴӢԜҢጄޣஒό৒ܰӢڙ၂ޣޑӸӧԶቹៜ ΑځՉࣁ߄౜Ƕ

--- ࿼კ΋ A ܭԜ --- (Β) U׎଎ࢭ

٬Ҕ U ׎଎ࢭޑЬाҞޑࢂҔٰෳ၂ڙ၂ޣჹܭ҂ٰஒ཮ᢀჸډ܈ࢂςᢀ ჸၸϕ୏ޑҢጄޣӧќ΋ঁόӕ൑ӦύϐϸᔈǶ଎ࢭЬा׷፦ࣁᓸլΚǴۭ೽ࣁ қՅǴନΑ଎ࢭύҔٰϩ႖όӕ୔ୱޑ౽୏Ԅ႖݈ࣁқՅѦǴ଎ࢭޑᕅᆶϺ޸݈

೿ࢂ೸ܴޑǴаճჴᡍޣᢀჸ୏ނޑࢲ୏ރݩǶӵკΒ A ܌ҢǴU ׎଎ࢭޑѰ ѓٿᆄϩձҗ΋ঁ V ࠠᖉ܌ಔԋǴ؂ঁ V ࠠᖉޑٿᖉ໔֨فࣁ 120±ȐٿᖉӚࣁ ߏ 42 Ϧϩǵቨ 10 Ϧϩǵଯ 15 ϦϩȑǴ؂ঁ V ࠠᖉഗᆄӚԖ΋ঁڈᐟނ୔Ǵҗ

(36)

қՅλ៓᠈ηȐߏ 8.5 Ϧϩǵቨ 8.5 Ϧϩǵଯ 13.5 Ϧϩȑ܌׎ԋǶӧ V ࠠٿᖉۭ

ᆄӚ࿼Ε΋Тё౽୏Ԅක݈Ȑό೸ܴԖऀϾޑᐋ✊࣒ዟ႖݈ǹࠆ 0.3 ϦϩǴϾ৩ ελࣁ 0.7 ϦϩȑǴԜٿТක݈ᆶ଎ࢭۭᜐޑڰۓԄᕅᏛȐό೸ܴԖऀϾޑᐋ✊

࣒ዟ႖݈ǴϾ৩ελࣁ 0.7 Ϧϩȑ׎ԋ΋҅Οف׎ޑ฻ࡑ୔ ȐᜐߏӚࣁ 10 Ϧ ϩȑǶӧ U ׎଎ࢭύǴ྽฻ࡑ୔ϐё౽୏Ԅක݈೏౽໒ਔǴନΑڈᐟނ୔ѦǴڙ ၂ޣёаᒧ᏷຾Ε଎ࢭޑҺ΋୔ୱǶࣁΑБߡ૶ᒵڙ၂ޣӧ U ׎଎ࢭύޑՏ࿼Ǵ

଎ࢭޑ V ࠠѰѓᖉӚёϩԋϖঁ୔ୱǴԶ V ࠠѰѓᖉё٩ᏵڈᐟނޑԖคϩձ

೏ڮӜࣁȨڀڈᐟނޑٗ΋ᖉȩᆶȨόڀڈᐟނޑٗ΋ᖉȩǴ؂ঁ V ࠠᖉΞёӆ ಒϩԋȨ᎞߈฻ࡑ୔ޑ฻ࡑ୔ȩᆶȨ᎞߈ڈᐟނޑڈᐟނ୔ȩǴ೭ٿঁ୔ୱΞё аਥᏵຯᚆᇻ߈ӆ຾΋؁೏֡ϩԋȨ߈฻ࡑ୔ȩȐߏ 21 ϦϩȑᆶȨᇻ฻ࡑ୔ȩȐߏ 21ϦϩȑаϷȨᇻڈᐟނ୔ȩȐߏ 21 ϦϩȑᆶȨ߈ڈᐟނ୔ȩȐߏ 21 Ϧϩȑǹନ ԜϐѦǴӧ߈ڈᐟނ୔ޑഗᆄຯᚆёаௗ᝻܈༘ᆪډڈᐟނ୔ 2 Ϧϩϣޑ୔ୱǴ

೏ᆀࣁȨ߈ڈᐟނ୔ޑന߻ᆄȩǶӢԜ U ׎଎ࢭύ؂ঁ V ࠠᖉϣޑ೭ϖঁ୔ୱவ ڈᐟނ୔ଆǴϩձࣁ߈ڈᐟނ୔ޑന߻ᆄǵ߈ڈᐟނ୔ǵᇻڈᐟނ୔ǵᇻ฻ࡑ୔ǵ аϷ߈฻ࡑ୔ǶࣁΑᙁϯჴᡍኧᏵБߡ౛ှଆـǴךॺஒȨ߈ڈᐟނ୔ޑന߻ᆄȩ у΢Ȩ߈ڈᐟނ୔ȩаϷȨᇻڈᐟނ୔ȩ೭Οঁ୔ୱӝᆀࣁȨښၰගٮ୔ȩǶ᏾

ঁ U ׎଎ࢭޑ೛ीࢂ௦ڗஏഈՠё೯॥ޑБԄǴ೸ၸܫ࿼ܭ଎ࢭۭᜐᕅᏛࡕБ ޑႝ୏ܜ॥৻Ǵܭჴᡍ຾Չύ໒௴Ǵ଎ࢭѦޑޜ਻ஒ࿶җڈᐟނ୔ܜΕٿᖉനࡕ ډၲ଎ࢭۭᜐࡕ௨рǶ

--- ࿼კΒ A ܭԜ --- ΟǵՉࣁբ཰ᆶࡼෳำׇ

ҁჴᡍ຾ՉޑՉࣁբ཰ࣁȨᢀჸᏢಞޑᏹ׋ȩǴҞޑࢂᡣ໢܄ঊႵѐᢀჸќ Ѧٿଫ໢܄ঊႵޑޗҬϕ୏ՉࣁǴᙖаࣴزޗ཮܄ૻ৲᠍ڗǶӧ໒ۈ຾ՉՉࣁჴ ᡍ߻Ǵջҗჴᡍޣ؂Ϻ׬ڗঊႵ٠ગख़ԿϿ΋ຼǴ٬ځಞᄍჴᡍޣޑᏹբǶՉࣁ

(37)

բ཰֡ӧ୏ނ܊਒ϐঊႵՉࣁෳ၂࠻ϣ຾ՉǴନΑϺ޸݈ޑ΋ЍआՅВӀᐩѦǴ ᗋ཮у΢຀௝๱ޑϤΜґᘚᐩ༾১ӀጕࣁྣܴȐྣܴࡋࣁ 30 LuxȑǶ܌ԖՉࣁᏹ

׋ᆶෳ၂֡Ծύϱ 12 ᗺၸࡕ໒ۈ຾ՉǴЪӧఁ΢Ύᗺа߻่״ǶӧԜբ཰ύǴ ڙ၂ޣܭᢀჸጃаϷ U ׎଎ࢭύޑ܌ԖՉࣁ߄౜ࣣаឪቹᐒȐJVCȑ૶ᒵΠٰǴ Զ୏ނޑ܌ԖՉࣁ߄౜֡аዸᒮ܈ឲᒵӧ฽૶ࠠႝတύޑՉࣁ૶ᒵ೬ᡏ Ethom

ٰ૶ᒵ (Shih & Mok, 2000)ǶӵკΟ܌ҢǴՉࣁჴᡍᆶᏹբำׇ΋Ӆೱុ຾Չ 6 ϺǴӧ೭ 6 Ϻύޑᏹ׋ёа٩ׇϩԋѤঁ໘ࢤǴϩձࣁಃ΋ϺޑȨಞᄍϯ໘ࢤȩ Ȑhabituation phaseȑǵಃΒϺޑȨ߻ෳ໘ࢤȩȐpretest phaseȑǵಃΟډಃϖϺ߻ъ ࢤޑȨᢀჸᏢಞ໘ࢤȩȐobservational learning phaseȑǵᆶಃϖϺࡕъࢤډಃϤϺ ޑȨ૶Ꮻෳ၂໘ࢤȩȐmemory testing phaseȑǶ೭Ѥঁ໘ࢤޑ၁ಒᏹ׋ำׇϩॊӵ ΠǺ

--- ࿼კΟܭԜ --- (΋) ಞᄍϯ໘ࢤ

Ԝ໘ࢤޑҞޑӧܭफ़եڙ၂ޣჹढ़ғᕉნޑขቾǴ٠٬ځዕ஼ՉࣁໆෳᏔ

׷ǶӧϤϺჴᡍࢬำޑಃ΋Ϻ཮຾Չಞᄍϯ໘ࢤǴ܌Ԗޑڙ၂ޣࣣ೏ϩձ࿼Ες

٬Ҕ 70 %ޑଚᆒᔔ࡬ଳృޑ U ׎଎ࢭύǴҺځԾҗ௖઩ 10 ϩដǶନΑ U ׎଎

ࢭѦǴڙ၂ޣࡕុΨ཮೏࿼ΕқՅλ៓᠈ηύǴܫΕᢀჸᏢಞጃϣޑᢀჸ୔຾Չ 30ϩដޑಞᄍϯǶ่״ࡕǴஒڙ၂ޣӼ࿼ӣচۚՐႵ᠈ϣǶ

(Β) ߻ෳ໘ࢤ

Ԝ໘ࢤޑҞޑࢂዴᇡஒ຾ΕᢀჸᏢಞ໘ࢤޑڙ၂ޣჹܭ҂ٰाᢀჸޑҢጄ ޣڀԖ΋ۓำࡋޑᑫ፪ǶӧϤϺჴᡍࢬำޑಃΒϺ཮຾Չ߻䬲໘ࢤǴ܌Ԗޑڙ၂ ޣࣣ೏ϩձ࿼Ες٬Ҕ 70 %ޑଚᆒᔔ࡬ଳృޑ U ׎଎ࢭύٰ຾Չෳ၂ǴԜ໘ࢤ х֖ٿঁ 3 ϩដޑೱុ჋၂ǴϩձࣁȨڙ၂ޣୃӳՏ࿼ෳ၂჋၂ȩᆶȨय़ჹڈᐟ ނޑᄊࡋෳ၂჋၂ȩǶӧڙ၂ޣୃӳՏ࿼ෳ၂჋၂ύǴ٠คڈᐟނޑӸӧǹय़ჹ

(38)

ڈᐟނޑᄊࡋෳ၂჋၂ύ܌٬ҔޑڈᐟނǴ߾ࢂڙ၂ޣ҂ٰӧᢀჸᏢಞ໘ࢤύ܌

࣮ډޑٿଫҢጄޣύޑځύ΋ଫǶڙ၂ޣୃӳՏ࿼ෳ၂჋၂ޑҞޑӧܭᒧрௗΠ

ٰा຾Չޑय़ჹڈᐟނޑᄊࡋෳ၂჋၂ύǴڈᐟނाܫ࿼ܭ U ׎଎ࢭޑѰᜐ܈

ѓᜐޑՏ࿼Ƕ໒ۈ຾ՉԜ჋၂ਔǴڙ၂ޣ཮೏࿼ܫܭ฻ࡑ୔ϣǴӧ॥৻໒௴ 30

ࣾࡕǴஒё౽୏Ԅක݈ܜ໒Ǵᡣڙ၂ޣӧ U ׎଎ࢭϣԾҗ௖઩Ƕҗܭ U ׎଎ࢭ Ѱѓ V ࠠᖉӚ೏Ⴤϩр 5 ঁ୔ୱǴӢԜ჋၂ύڙ၂ޣӧ೭Ӆ 10 ୔ޑӚ୔ୱϣϐ

೰੮ਔ໔аϷӚ୔ୱϐऀຫԛኧ཮೏ჴᡍޣ૶ᒵΠٰǴڙ၂ޣӧӚ୔ୱޑऀຫԛ ኧຎࣁڙ၂ޣӧӚ୔ୱϐࢲ୏Κޑࡰ኱ǶӧԜ჋၂΢܌ໆෳޑڙ၂ޣϐՉࣁࡰ኱

ࣁȨU ׎଎ࢭϐࢲ୏ΚȩǴջڙ၂ޣӧ U ׎଎ࢭϣޑ 10 ঁ୔ୱ໔ޑᕴऀຫԛኧǶ

྽ 3 ϩដޑ௖઩ਔ໔ډၲਔǴஒ॥৻ᜢ௞Ǵڙ၂ޣ߾೏ܫӣ฻ࡑ୔ύǴ฻ࡑΠ΋

ঁय़ჹڈᐟނޑᄊࡋෳ၂჋၂Ƕ

ڙ၂ޣӧय़ჹڈᐟނޑᄊࡋෳ၂჋၂΢܌ाय़ჹޑڈᐟނࢂ௦ڗᒿᐒϩ ࢴǴӧڈᐟނޑᒧ᏷΢ǴࣁΑ௓ڋڙ၂ޣ҂ٰ܌ᢀჸډҢጄޣ۶Ԝ໔ޑѺରࠔ፦

ᆶയॄ௃׎Ǵךॺ٣Ӄ૽ግ٠ࡷᒧр΋ץڀԖೱՍᕇയ࿶ᡍޑគৎЪၨڀமװᔐ

܄ޑ΋ץװᔐޣȐfighterȑǴ஑ߐҔٰѺॹჹЋǹӕਔΨྗഢΑ΋ץڀԖऩυѨ

௳࿶ᡍЪၨόڀװᔐૈΚޑᒡৎǴ٬ځӧय़ჹڀװᔐ܄ޑჹЋ೯தѝԖ਌Ѻޑ ҽǶӧ߻ෳ໘ࢤ໒ۈ຾Չ߻Ǵךॺ཮ᙣ཈ޑᡣӚঁҢጄޣ೿ᆶځѬӕᜪ຾Չၸӭ ԛޑޗҬϕ୏Ǵٰዴߥځ܌ڀيҽᗋࢂᆢ࡭όᡂޑǶय़ჹڈᐟނޑᄊࡋෳ၂჋၂ ޑᏹ׋ำׇӵӕڙ၂ޣୃӳՏ࿼ෳ၂჋၂΋ኬǴڈᐟނ܌ᘍܫޑՏ࿼ࢂҗڙ၂ޣ ӧ຾Չڙ၂ޣୃӳՏ࿼ෳ၂჋၂ਔǴࡑӧ U ׎଎ࢭٿᖉύځύ΋ᖉϣຯᚆڈᐟ ނ୔ന᎞߈ϐȨ߈ڈᐟނ୔ޑന߻ᆄȩу΢Ȩ߈ڈᐟނ୔ȩ೭ٿ୔ޑᕴਔ໔ၨߏ ޣࣁڈᐟނܫ࿼ޑ΋ᆄǶӕኬ૶ᒵ჋၂ύڙ၂ޣӧӚ୔ୱϐऀຫԛኧᆶ೰੮ਔ ໔ǶӧԜ჋၂΢ाໆෳޑڙ၂ޣϐՉࣁࡰ኱ӅԖѤঁǴϩձࣁڀڈᐟނޑٗ΋ᖉ ϐȨډၲ߈ڈᐟނ୔ޑന߻ᆄϐ܌ሡਔ໔ȩᆶȨښၰගٮ୔ϐ೰੮ਔ໔ȩǹȨόڀ ڈᐟނޑٗ΋ᖉϐࢲ୏ΚȩǵᆶȨόڀڈᐟނޑٗ΋ᖉϐ೰੮ਔ໔ȩǶ྽჋၂่״

ࡕǴஒڙ၂ޣӼ࿼ӣচۚՐႵ᠈ϣǴU ׎଎ࢭ߾٬Ҕ 70 %ޑଚᆒᔔ࡬ଳృᗉխ

(39)

੮ΠҺՖ਻ښǶ

ࣁΑዴᇡڙ၂ޣჹڈᐟނڀԖ΋ۓำࡋޑᑫ፪ᆶݙཀำࡋǴךॺۓΠΑᑔᒧ

኱ྗࣁǺӧय़ჹڈᐟނޑᄊࡋෳ၂჋၂ޑ 3 ϩដϣǴѸ໪೰੮ӧڀڈᐟނޑٗ΋

ᖉຯᚆڈᐟނ୔ന᎞߈ϐȨ߈ڈᐟނ୔ޑന߻ᆄȩу΢Ȩ߈ڈᐟނ୔ȩ೭ٿঁ୔

ୱޑਔ໔ᕴکεܭ 60 ࣾа΢Ǵ܈ޣࡑӧȨ߈ڈᐟނ୔ޑന߻ᆄȩޑਔ໔Ԗ 40

ࣾа΢Ƕ೯ၸ߻ෳ໘ࢤᑔᒧ኱ྗޑڙ၂ޣஒ೏ᒿᐒϩࢴډΟಔǴϩձࣁȨय़ჹύ

܄ڈᐟಔȩǵȨय़ჹᒡৎڈᐟಔȩǵᆶȨय़ჹគৎڈᐟಔȩǶӧֹԋ U ׎଎ࢭޑ߻

ෳࡕǴ೭٤ڙ၂ޣӆ΋ԛ೏࿼ܫΕᢀჸᏢಞጃϣޑᢀჸ୔຾Չ 30 ϩដޑಞᄍ ϯǴ٬ځкϩ௖઩٠ಞᄍᢀჸ୔аϷλ៓᠈Ƕ

(Ο) ᢀჸᏢಞ໘ࢤ

Ԝ໘ࢤޑҞޑࢂᡣڙ၂ޣೱុΟϺᢀჸځѬٿଫ໢܄ঊႵวғޗҬϕ୏٣ ҹǴ٠೸ၸᢀჸ೭ኬޑϕ୏٣ҹٰᏢಞ୔ϩ܌ᢀჸډޑҢጄޣϐ໔ޑόӕǶӧϤ ϺჴᡍࢬำޑಃΟϺ཮໒ۈ຾ՉೱុΟϺޑᢀჸᏢಞ໘ࢤǴڙ၂ޣ೏ᒿᐒϩࣁჴ ᡍಔᆶ௓ڋಔǴჴᡍಔ཮ᢀჸډٿଫ໢܄ঊႵޔௗϕ୏٠วғѺର٣ҹǴᆀϐࣁ ȨᢀჸѺର٣ҹಔȩǹ௓ڋಔ߾ᢀჸٿଫ໢܄ঊႵ႖๱λ៓᠈໔ௗϕ୏ՠคݤว ғѺର٣ҹǴᆀϐࣁȨᢀჸύ܄٣ҹಔȩǶӵკѤ܌ҢǴӧᢀჸύ܄٣ҹಔύǴ ךॺஒځύ΋ଫҢጄޣᜢΕλ៓᠈ηύǴ࿼ܭϕ୏୔ޑύѧǴԶᡣќѦ΋ଫҢጄ ޣࡑӧϕ୏୔ύޑ៓᠈ηѦय़Ǵ٬ځᆶ៓᠈ηϣޑҢጄޣ຾Չ໔ௗ܄ϕ࣬௖઩ޑ ՉࣁՠόวғѺରՉࣁǶӃ߻೯ၸ߻ෳ໘ࢤᑔᒧޑڙ၂ޣǴ཮ਥᏵځӧ၀໘ࢤਔ

܌य़ჹޑڈᐟނϐيҽङඳǴ຾Զஒڙ၂ޣᒿᐒϩࢴډᢀჸѺର٣ҹಔ܈ᢀჸύ

܄٣ҹಔǶКӵӧ߻ෳ໘ࢤਔǴӧჴᡍޣޑᒿᐒϩࢴΠǴӵ݀ڙ၂ޣय़ჹޑڈᐟ ނޑيҽङඳࣁᒡৎޑҢጄޣǴ߾ᢀჸᏢಞ໘ࢤਔǴڙ၂ޣ൩཮೏ϩࢴډᢀჸѺ ର٣ҹಔύǴӵԜڙ၂ޣωૈᢀჸډيҽङඳࢂᒡৎޑҢጄޣ೏ќ΋ଫគৎҢጄ ޣ๏Ѻ௳ǹӵ݀ڙ၂ޣय़ჹޑڈᐟނيҽङඳࢂគৎǴ߾ᢀჸᏢಞ໘ࢤਔǴڙ၂ ޣஒᒿᐒ೏ϩࢴډᢀჸύ܄٣ҹಔ܈ޣᢀჸѺର٣ҹಔϐύǴӵԜڙ၂ޣஒ཮ᢀ ჸډԜڈᐟނيҽङඳࣁគৎޑҢጄޣǴ཮ܭύ܄٣ҹύᆶќ΋ଫҢጄޣ຾Չύ

(40)

܄ϕ୏Ǵ܈ܭѺର٣ҹύѺ௳ќ΋ଫᒡৎҢጄޣǶӢԜǴय़ჹύ܄ڈᐟಔȐn = 13ȑ ޑڙ၂ޣᢀჸޑࢂٿଫҢጄޣ຾Չύ܄ϕ୏Ǵջࣁᢀჸύ܄٣ҹಔȐn = 13ȑǹԶ य़ჹᒡৎڈᐟಔȐn = 10ȑᆶय़ჹគৎڈᐟಔȐn = 11ȑύޑڙ၂ޣӧΟϺޑᢀჸ Ꮲಞ໘ࢤӕኬᢀჸٿଫҢጄޣ຾ՉѺରϕ୏ǴջࣁᢀჸѺର٣ҹಔȐn = 21ȑǶ

--- ࿼კѤܭԜ ---

ಃΟϺ཮຾ՉᢀჸᏢಞ໘ࢤޑಃ΋Ϻ૽ግǴӧ૽ግ໒ۈ߻཮ஒό೸ܴό೸਻

ޑᓸլΚ݈ߔ႖ӧᢀჸ୔ᆶϕ୏୔ϐ໔Ǵ٬ڙ၂ޣӧ຾Εᢀჸ୔ࡕኩਔ࣮όډϕ

୏୔ϣޑҺՖࢲ୏Ƕ྽ڙ၂ޣ೏࿼ܭᢀჸ୔ࡕǴٿଫջஒा຾Չϕ୏ޑҢጄޣޑ ځύ΋ଫஒӃ೏࿼Εϕ୏୔ύǴԶќ΋ଫҢጄޣ߾೏ܫΕ฻ࡑ୔ύ฻ࡑǶ྽॥৻

໒௴ࡕǴᢀჸ୔کϕ୏୔໔аϷϕ୏୔ک฻ࡑ୔໔ޑ႖݈཮೏ӕਔܜ໒Ǵ྽฻ࡑ

୔ύޑҢጄޣ຾Εϕ୏୔ࡕǴ൩ஒϕ୏୔ᆶ฻ࡑ୔໔ޑ႖݈ܫӣǶڙ၂ޣࡑӧᢀ ჸ୔ޑᢀჸਔ໔཮٩Ңጄޣޑ߄౜ԶۓǴҗჴᡍޣ૶ᒵҢጄޣ۶Ԝޑϕ୏௃ݩǴ

྽Ңጄޣ۶ԜԖϕ୏ਔ཮Ҕዸᒮ຾Չीਔǹϕ୏ଶЗਔ߾ኩଶीਔǴ྽Ңጄޣ۶ ԜԖϕ୏Զਔ໔ಕᑈԿ 1 ϩ 30 ࣾ܈ޣ྽ٿଫҢጄޣࡑӧϕ୏୔ϣޑᕴਔ໔ၲډ 29ϩដংǴջଶЗಕᑈਔ໔ǹ1 ϩដࡕǴ྽ϺޑᢀჸᏢಞ૽ግ൩೏ಖЗǶᢀჸѺ ର٣ҹಔޑҢጄޣϕ୏ۓကࣁǺيҽङඳࣁគৎޑҢጄޣ஭቏టࠈќ΋ଫيҽङ ඳࣁᒡৎޑҢጄޣǴՠόхࡴគৎޑҢጄޣ೏ᒡৎޑҢጄޣᓸॹӧӦޑ஭቏టϸ ᔐǴନߚϸᔐԋфωᆉǹᢀჸύ܄٣ҹಔޑҢጄޣϕ୏ۓက߾ࣁǺλ៓᠈ηѦޑ Ңጄޣჹ៓᠈ηϣޑҢጄޣ຾Չޔௗޑ༘ᆪ୏բǶ྽ᢀჸᏢಞ૽ግ่״ਔǴ॥৻

ᜢ௞ǴқՅό೸਻႖݈Ψӆ࿼ӣᢀჸ୔ᆶϕ୏୔ϐ໔ǴฅࡕஒҢጄޣڗрǵӆڗ рڙ၂ޣǴ࿼ӣচۚՐϐႵ᠈ϣǶᢀჸᏢಞ໘ࢤޑಃΒᆶಃΟϺ૽ግǴ܌Ԗჴᡍ ำׇᆶᢀჸᏢಞ໘ࢤಃ΋Ϻ૽ግࢂֹӄ࣬ӕޑǶࣁΑዴᇡڙ၂ޣӧ૽ግಃ΋Ϻа Ϸനࡕ΋ϺޑѺର٣ҹύ܌ᢀჸډޑҢጄޣޑ߄౜όࢂ೷ԋࡕុޑ૶Ꮻෳ၂໘ ࢤ΢όӕಔڙ၂ޣϐϸᔈԖ܌ৡ౦ޑӢનǴӢԜךॺа૽ግಃ΋ϺаϷಃΟϺ܌

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