୯ҥᆵεᏢᏢଣЈᏢࣴز܌
ᅺγፕЎ
Graduate Institute of Psychology College of Science
National Taiwan University Master Thesis
ঊႵΨՉޗ܄ૻ৲᠍ڗЪЇวᢀჸᏢಞΠ!
аϷ٣ࡕޑׯᡂ !
Hamsters Eavesdrop Too and The Consequent Changes during and after Observational Learning
ܴے Ming-Tsung Kuo
ࡰᏤ௲ǺᒘЎ റγ Advisor: Wen-Sung Lai, Ph.D.
ύ҇୯ 98 ԃ 7 Д
July, 2009
ᇞᖴ
ӳǼᕴᆉֹԋ೭ҁፕЎΑǴܴےǴ༸ளӳǼӃ๏ԾρٰঁඓᖂփǴǼǼ
ǼǼΟԃΑǴӣགྷ߃ǴϙሶόᔉǴ൩೭ኬᅈ๚Ո፯Ε೭ঁࣽᏢᘀ
݅ǴΟԃၸѐΑǴᗨฅၹᡏᡒǴՠЈǴ׳уय֧ǶाགᖴޑΓϼӭǴόѝࢂᖴ ϺǶགᖴךޑР҆ᆶঢঢکۆۆǴؒԖգॺǴόԖϞϺޑךᆶךޑܴϺǶךޑ
ࡰᏤ௲ǴᒘЎԴৣǴᖴᖴா೭Οԃٰޑаيբ߾ᆶ᠋ǴᖴᖴǼགᖴలζԶ ۆǴךࣴز܌ғఱύޑΓǴᖴᖴாޔаٰעךٽηྣ៝ǴᖴᖴǶఉٍ۪
ԴৣǴᖴᖴாǴӵ݀ϖԃؒԖ᠋ၸாளፐǴךёૈᗋόᏢคೌǴόޕՖᒏғ
ЈᏢǴ׳ᖴᖴாѳВޑᜢЈǶسޑߋӃғǴᖴᖴாதӧךႝတрୢᚒਔр Ћ࣬շǴϼགᖴΑǼᖴᖴᗛቺᏦԴৣᆶߋඵ፣Դৣӧךޑჴᡍ๏ϒڐշᆶࡰ
ᏤǴᗨฅ࣬ೀਔ໔όߏǴՠߚதᖴᖴǶᖴᖴ৪ֻԴৣᆶᎄᖚԴৣᜫཀךޑ α၂ہ٠๏ϒᝊޑࡌǶғ܌ޑࡼ࣓࣓ᏢۆᆶεᏢǴᖴᖴգॺӧךޑ ჴᡍ๏ϒڐշᆶࡰᏤǴᖴᖴǼሺོǵৎྍǵЎᇬǵࣁฺǵӵదǵаϷॕ⪭Ǵག ᖴգॺჴᡍޑڐշǶΓيፏԵᡍǴόࢂाᔐॹգ Զࢂाԋ൩գǴࠂᏌᏢߏǵ
܃ЎǵᝬЬǵаϷ㾈ሺǴғڮύԖգॺޑрࢂךࣴز܌ޑВη္Ǵԋߏࡐε ޑΚٰྍǴᖴᖴǼШၲᏢߏǵើᏢۆǵаϷไᏢۆǴᖴᖴգॺӧךԖሡा
ਔεΚ࣬շǶζΐᓓޑӚՏߓ࠰ॺǴߚதགᖴգॺ೭Οԃჹךޑྣ៝Ǵத๏ך уǴϼགᖴǼགᖴך܌Ԗޑӳ϶ॺǴᖴᖴգॺૈхךǴ҂ٰᗋፎӭӭх֖Ƕ നࡕǴךाߚதᖴᖴமԶԖΚམӧךުޑออǴךന഻یΑǼЋЋǴଆ
փǶ
ᄔा
ᢀჸᏢಞǴཀࡰғЋၸᢀჸдΓޑՉԶಞளཥޑՉࣁ܈ૻ৲Ǵ೭ᅿᏢ ಞБԄςӧόӕނᅿޑፏӭՉࣁύჴၸǶၸᢀჸдΓޑޗϕԶவύ
ڗрૻ৲ޑ೭ᅿᏢಞБԄᆀࣁޗ܄ૻ৲᠍ڗǹၸѐ࣬ᜢޑࣴزεӭଞჹΓ ᜪаϷࢌ٤ങᜪᆶചᜪǴՠଞჹΓᜪаѦޑথ٢ᜪނᗋ҂මԖၸǶճҔ௶ճ٥
܄ঊႵڀԖϐሦୱװᔐՉࣁǴךॺวрঁՉࣁᏹբБݤٰޗ܄ૻ৲
᠍ڗǶҁፕЎύӅԖΟঁЬჴᡍǴҞޑӧᢀჸޣೱុΟϺჹٿঁҢጄޣޑѺ ର܈ύ܄٣ҹՉᢀჸਔ܌ЇวᢀჸᏢಞΠаϷ٣ࡕϐՉࣁᆶᓸΚລᅟᆾޑ ׯᡂǶӧჴᡍύǴ҂ԖѺରᡍޑ܄ᢀჸޣǴӧΟϺᢀჸ่״ࡕҥջ܈ࢂ
ϺࡕǴܭ U ࢭϣזೲޑᖿ߈٠ᡉӦၨӭਔ໔ѐគৎҢጄޣǴᡉ Ң܄ᢀჸޣКၨԖᑫ፪ᆶគৎҢጄޣՉϕǶ࣬ၨܭჴᡍύ٬ҔΑ҂ԖѺ ରᡍϐᢀჸޣǴӧჴᡍΒύ߾ࢂ٬ҔΑՉᢀჸᏢಞϺමᎁၶढ़ғװᔐޣ
௳ၸԛޑ܄ᢀჸޣǴ٠ว೭٤ᢀჸޣӧᐕೱុΟϺᢀჸѺର٣ҹޑ
ΠаϷࡕុӧ U ࢭύޑҥջکߏයෳ၂Ǵ֡߄рᆶঁჴᡍֹӄ
࣬ϸޑՉࣁኳԄȐх֖Ǻၨӭޑᕨߡ௨ܫǵၨӭޑາᆶᗉǵаϷၨϿޑ
ਔ໔ȑǴՠԜਏ݀٠όрӧೱុΟϺᢀჸύ܄٣ҹϐԖ௳ᡍޑᢀჸޣ
Ƕ೭٤่݀ᡉҢΑ௳ᡍૈԖਏӦቹៜڙঊႵځࡕޑՉࣁ߄ǶӧჴᡍΟ ύǴ٬ҔΑߚߟΕ܄ޑБݤٰୀෳᢀჸޣᕨߡϣҜ፦ᎇжᖴނȐFCMȑޑ֖ໆǶ
ӧჴᡍΟ A ύǴၸ 3 ঁλਔᇆԛᕨߡኬҁޑ௦ኬБݤǴࣴز่݀ᡉҢ ӧᐕԛװᔐϕࡕǴᒡৎᕨߡϣޑҜ፦ᎇжᖴނᐚࡋܭ٣ҹࡕޑಃΟᆶಃΒ
ΜλਔԖܴᡉޑϲǴՠ೭ኬޑׯᡂ٠όวғӧគৎيǶӧჴᡍΟ B ύǴ߾٬ҔΑ٩ᏵჴᡍΟ A ่݀অׯၸޑ 18 λਔᕨߡ௦ݤٰϩჴᡍΒύ ޑᢀჸޣ܌௨ܫрޑᕨߡኬҁǶӧΟϺޑᢀჸύǴԖ௳ᡍޑ܄ᢀჸޣӧᢀ ჸѺର٣ҹਔǴځ௨ᕨໆԖܴᡉޑቚуǴՠځᕨߡϣҜ፦ᎇжᖴނޑᐚࡋࠅ࣬ჹ
եܭᢀჸύ܄ϕ٣ҹޑᢀჸޣǶӕኬӦӧ่״ᢀჸࡕϺǴय़ჹគৎҢጄޣ߄
ᐟਗ਼ޑᢀჸޣيΨុໆෳрᕨߡϣҜ፦ᎇжᖴނफ़եޑǶᆕӝа܌
ॊǴ೭٤่݀ᡉҢΑ܄ঊႵΨૈՉޗ܄ૻ৲᠍ڗǴЪঁᡏၸѐޗҬ௳ޑ
ᡍᡉӦቹៜځࡕុၶډ೭٤ወӧჹЋਔૻ৲ޑ٬ҔаϷຑޑБԄǶᙖҗ ᢀჸ܌ᕇளٰޑૻ৲ǴόѝቹៜΑғޑࡡۓ܄ǴΨቹៜᒿࡕޑՉࣁϸᔈǶ
ᣂڗΚᨠኘᖂΕषᄎࢤಛஒ࠷ΕᚘԺ֘ᚨΕސᚰࢤΕᜈঁփؼᔆᔧזढΕ ପቕΖ
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
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.
Ҟԛ!
!
ಃകʳ ᆣፕ////////////////////////////////////////////////////////1!
ಃʳ Їق////////////////////////////////////////////////////1 ᢀჸᏢಞ////////////////////////////////////////////////////1
ಃΒʳ ӣ៝ϞӸӧܭނيޑᢀჸᏢಞࣴز////////////////////3
൘ǵᆶғӸԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////////3 ມǵᆶ܄ᆶᕷ़ԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////6
ୖǵᆶٛᑇԖᜢޑᢀჸᏢಞՉࣁ////////////////////////////////7 စǵኳү////////////////////////////////////////////////////9
ಃΟʳ ޗ܄ૻ৲᠍ڗ/////////////////////////////////////////12
൘ǵวғܭޗ܄ᝡݾύޑޔௗຑᆶ໔ௗຑ/////////////////12 ມǵޗ܄ᝡݾύଞჹޗ܄ૻ৲᠍ڗӧނيޑࣴز/////////14
ୖǵঁᡏᡍჹޗᝡݾύޑޗ܄ૻ৲᠍ڗޑቹៜ/////////////17
ಃѤʳ ҁፕЎޑࣴزᚒᆶౣ/////////////////////////////////18
൘ǵࣴزᚒ///////////////////////////////////////////////18 ມǵਡЈୢᚒᆶှ،ౣ/////////////////////////////////////22
ಃΒകʳ ჴᡍᆶБݤ/////////////////////////////////////////////25!
ಃʳ ჴᡍ/////////////////////////////////////////////////25
ಃΒʳ ჴᡍΒ/////////////////////////////////////////////////33
ಃΟʳ ჴᡍΟ/////////////////////////////////////////////////36
൘ǵჴᡍΟ A///////////////////////////////////////////////37 ມǵჴᡍΟ B///////////////////////////////////////////////41
ಃΟകʳ ่݀ᆶፕ/////////////////////////////////////////////////45!
ಃʳ ჴᡍ/////////////////////////////////////////////////45
҂ԖޗҬᡍޑᢀჸޣӧޗ܄ૻ৲᠍ڗޑᐕำύᆶࡕុૻ৲٬
ҔਔޑՉࣁ߄/////////////////////////////////////////////45
ಃΒʳ ჴᡍΒ/////////////////////////////////////////////////52
ڀԖԛޗҬ௳ᡍޑᢀჸޣӧޗ܄ૻ৲᠍ڗޑᐕำύᆶࡕ
ុૻ৲٬ҔਔޑՉࣁ߄/////////////////////////////////////52
ಃΟʳ ჴᡍΟ A///////////////////////////////////////////////62
ԛޗҬ௳ᡍΠޑᓸΚϸᔈ///////////////////////////62
ಃѤʳ ჴᡍΟ B///////////////////////////////////////////////67
ڀԖԛޗҬ௳ᡍޑᢀჸޣӧޗ܄ૻ৲᠍ڗޑᐕำύᆶࡕ
ុޑૻ৲٬ҔਔޑᓸΚϸᔈ///////////////////////////////////67
ಃѤകʳ ᆕӝፕ///////////////////////////////////////////////////73!
ಃʳ ճҔ܄௶ճ٥ঊႵނኳԄࣴز࣬ᜢᚒޑᓬ༈///////////74
ಃΒʳ ҂ԖޗҬᡍΠޑእЋຑᐕำ///////////////////////////77
ಃΟʳ ڀԖޗҬ௳ᡍΠޑእЋຑᐕำ///////////////////////78
ಃѤʳ ঁᡏޗҬ௳ᡍჹޗ܄ૻ৲᠍ڗᐕำᆶࡕុޑእЋຑᐕำ ϐቹៜ܄///////////////////////////////////////////////79
ಃϖʳ ޗ܄ૻ৲ௗڙਔᆶঁᡏӆᇡਔ܌ճҔޑጕ///////////////81
ಃϤʳ ᓸΚᡍΠޑঁᡏৡ౦܄/////////////////////////////////83
ಃΎʳ ҁፕЎޑଅᆶځ҂ٰޑᔈҔ/////////////////////////////85
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ߕკᇥܴ///////////////////////////////////////////////////////////95!
ߕკ//////////////////////////////////////////////////////////////103!
ߕ߄//////////////////////////////////////////////////////////////117!
ߕᒵ//////////////////////////////////////////////////////////////127
ಃ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
ᏢಞǴջނᅿࣁΑૈᔈᕉნזೲᡂϯ܌ᄽϯрޑᅿૈΚǴѬ٩ᕉნޑ ᡂϯԶрޑፓǴ܌٩ᒘޑջࣁၸѐჹ၀ڈᐟ܈၀ᕉნޑᏫǶᗨฅӵ ԜǴᏢಞޣ۳۳ሡाᒃԾѐᐕ٠бржሽǴωளаᕇளᡍǴٯӵǺၸᒃԾ ӞΠࢌނᡏࡕᏤठғੰǴωΑှډԜނᡏࢂόё१ޑǶ
ᢀჸᏢಞ
ঁᡏᏢಞȐindividual learningȑ೯தሡाၸޔௗѐᐕ٣ҹωૈᏢಞډǴ Ψ۳۳ӢԜбрΑۓޑжሽǹ࣬ჹԶقǴќᅿ໔ௗԄޑᏢಞ߾ࢂၸᢀჸځ дΓޑᡍǴ൩ૈڙډቹៜ܈ᕇளૻ৲Ǵ೭൩ࢂ܌ᒏޑᢀჸᏢಞȐobservational learningȑǶӧঁᡏᏢಞᐕำύǴ೯தሡाᐕۓำࡋޑ၂ᇤᆶ၂Ǵωૈගଯ ಞள၀Չࣁޑёૈ܄ǹ࣬ჹԶقǴᢀჸᏢಞѝሡाၸᢀჸձΓޑՉࣁ߄Ǵ൩
ૈගଯಞள၀Չࣁޑёૈ܄ǴӢԜᢀჸᏢಞёаεεफ़ե܌ሡбрޑжሽǶᢀჸ ᏢಞԖځख़ाޑғӸሽॶǴΨ֎Ї٤ࣽᏢৎΕ೭ঁሦୱޑࣴزǶّϞቺ
ȐAlbert Banduraȑӧ 1977 ԃ܌ගрޑȨޗᏢಞፕȩȐSocial learning theoryȑǴ ගٮΑঁࣴزޑख़ाୖԵаϷፕ୷ᘵǶ
ӧቺޑޗᏢಞፕύǴჹՉࣁޑಞள௦ڗ׳ቶݱ܄ޑᢀᗺȐBandura;
1977/1995ȑǴᙁൂٰᇥǴቺᇡࣁՉࣁޑಞளࡽڙᒪӢનޑज़ڋǴΞڙࡕϺ
ᡍᕉნޑቹៜǴࡐᜤஒٿޣϩ໒ǶΓޑፄᚇՉࣁЬाࢂ٩ᒘࡕϺಞளޑǴԶᇡ
ޕתᄽΑঁࡐख़ाޑӢનǶՉࣁಞளԖٿᅿόӕޑၸำǴᅿࢂ೯ၸޔௗ
ᡍǴᕇளՉࣁᆶϸᔈ่݀۶Ԝ໔ᜢ߯ޑၸำǴᆀࣁȨ೯ၸϸᔈޑ่݀܌ՉޑᏢ ಞȩǹќᅿࢂӧޗნΠǴᢀჸޣȐobserverȑ೯ၸȸᢀჸȹҢጄޣ
ȐdemonstratorȑޑՉࣁԶಞளՉࣁޑၸำǴᆀࣁȨ೯ၸҢጄޣ܌ՉޑᏢಞȩǶ
ಃᅿՉࣁᕇளБԄջࣁঁᡏᏢಞǴӧ೭ᅿБԄǴϸᔈޑ่݀ϐ܌аჹᏢಞࢂ
ԖᔅշޑǴЬाࢂӢࣁѬගٮΑΟᅿЬाфૈǴϩձࢂૻ৲ǵᐒǵᆶቚமфૈǶ ӧૻ৲фૈǴঁᡏૈவՉࣁϸᔈᆶ၀Չࣁ܌ٰޑ่݀ԶᕇளૈࡰᏤΠԛՉ
ޑૻ৲ǹӧᐒфૈǴӃޑᡍ٬ளঁᡏჹܭࢌՉޑ่݀ૈԖႣޕޑ གྷႽǴԶ٬ளঁᡏѐᒧՉ܈όՉ၀ՉǹӧቚமфૈǴΑှՖᅿՉ
ૈٰՖᅿၿ፞Ǵჹܭ၀Չࢂցૈߦ܈ڋࢂख़ाޑǶಃΒᅿՉࣁᕇளБ ԄջࣁᢀჸᏢಞǴӧ೭ᅿБԄǴૻ৲ǵᐒǵᆶቚம೭٤фૈ߾ૈӧঁᡏόሡ ᒃيѐՉ၀ՉջૈၸᢀჸձΓޑՉ൩ૈᕇளǴΨӢԜ೭ኬޑᏢಞБԄǴ ჹΓᜪεӭኧޑՉࣁᕇளࢂख़ाޑǶቺᇡࣁǴᢀჸᏢಞࣁೱុޑᐕำǴԖ ѤঁЬाᐕำǴϩձࣁݙཀᐕำǵߥ੮ᐕำǵՉࣁᇙᐕำǵϷᐒᐕำǶݙཀ ᐕำࢂᢀჸᏢಞวғޑಃǴӧݙཀᐕำύǴҢጄޣՉҁيޑቻ܈மࡋǵ ᢀჸޣၸѐޑᡍǵаϷᢀჸޣکҢጄޣϐ໔ޑᜢ߯ፏӭӢનቹៜᏢಞޑ ਏ݀ǶӧᢀჸᏢಞޑߥ੮ᐕำǴᢀჸޣૈஒ܌ᢀჸډޑ٣ҹᙯϯࣁࢌᅿ಄ဦࠠ
ԄǴջ٬ҢጄޣόӆрǴΨૈၸ၀಄ဦࣁ൞ϟԶஒ܌ᢀჸډޑૻ৲੮ӧอය
Ꮻ܈ߏයᏫύǶӧՉࣁᇙᐕำǴᢀჸޣሡाעᏫύޑ಄ဦᙯඤԋޑ ՉࣁǴ೭ၸำੋϷډسӈᇡޕޑکՉࣁޑᏹբǴКӵჹՉࣁӵՖ߄ޑᇡ ޕǹૈჸԾيՉᆶᏫύҢጄޣޑՉϐ໔ޑৡ౦٠Չঅ҅Ƕӧᐒᐕำ
Ǵᢀჸޣჹܭ܌ᢀჸޑ٣ҹࢂցڀȨයࡑȩǴ܈ޣाᢀჸޣѐ߄ޑნࢂ
ցӝܭᢀჸޣӧޑރᄊǴࣣቹៜᢀჸޣࢂցૈத߄р܌ಞளޑՉ ࣁǶ೭ѤঁᐕำޑൻᕉՉǴ٬ளനಖՉࣁૈ٩ᒘᢀჸᏢಞޑБԄԶᕇளǶ
ቺ 1977 ԃޑޗᏢಞፕǴаȨᇡޕȩޑཷۺٰඔॊΓᜪޑՉࣁᕇளǴ ԶΓᜪޑεϩՉࣁёаᙖҗᢀჸᏢಞ೭ኬޑБԄٰᕇளǶӵ݀ाჹΓᜪޑՉࣁ
ᕇளᆶځङࡕޑЈᐒڋԖ׳ӭޑΑှǴךॺ൩ѸाჹᢀჸᏢಞ೭ኬޑᏢಞБ ԄԖ׳ӭޑࣴزᆶǶҗܭတઓࣽᏢӧ߈жޑזೲஙଆǴೱӦߦԋޗઓ
ࣽᏢޑᑫଆǴӢԜȨᢀჸᏢಞȩङࡕޑઓᐒڋᅌԋࣁ߈жઓࣽᏢৎॺո ΚགྷѐᑍޑߐᚒǴՠҗܭаΓᜪࣁڙ၂ޣததԖፏӭज़ڋǴा׳ుΕӦѐ
ϣӧޑЈᐒڋࣗԿܭઓᐒڋࢂόܰޑǶ࣬ၨϐΠǴނኳࠠගٮΑߡ ճ܄ǵܰᏹբ܄ǵаϷӢ݀ፕ࣬ჹᙁൂӳೀǴӢԜӵ݀ૈၸނኳࠠ
ٰࣴزᢀჸᏢಞ೭ኬޑᚒǴόԖշܭ҂ٰᔈҔډΓᜪޑᢀჸᏢಞǴΨԖշܭ ךॺѐΑှނޑᇡޕૈΚǶ
ಃΒʳ ӣ៝ϞӸӧܭނيޑᢀჸᏢಞࣴز
ၸѐଞჹόӕނᅿᆶόӕՉࣁԖፏӭࣴزҾკѐᢀჸᏢಞޑᚒǴԶ ёૈቹៜᢀჸᏢಞޑӢનΨԖӣ៝Ўക၁ಒӦፕ (Zentall, 2006)Ƕၸӣ៝ᆶ
ፕ೭٤ࣴزǴךॺёаᏢಞډӵՖीঁ࣬ჹֹޑჴᡍѐࣴزวғܭނ يޑȸᢀჸȹᆶȸᏢಞȹǶ
൘ǵᆶғӸԖᜢޑᢀჸᏢಞՉࣁ
ǵᏢಞ၀Ӟϙሶ१ނ
ଞჹ೭ᜪޑࣴزᚒǴӭъࢂࣴز٩ᒘ༘ޑނࣁЬǴനத٬Ҕޑނኳ
ࠠࢂȨαښୃӳޑޗሀբȩȐsocial transmission of flavor preferences paradigmȑǶ၀բޑ୷ҁჴᡍำׇࣁǺ૽ግঁҢጄޣѐ१ҔӃவؒӞၸޑ ढ़ғ१ނࡕǴᡣҢጄޣᆶќᢀჸޣ࣬ೀࢤਔ໔ǴӆஒҢጄޣڗр٠ᡣᢀჸޣ ჹٿᅿढ़ғ१ނٰՉᒧǶᢀჸޣӧٿᅿढ़ғ१ނޑᒧӛᒧҢጄޣӞ ၸޑ१ނǴ೭ኬޑ่ܴ݀ᢀჸޣӧढ़ғޑރݩΠӛճҔΑΓޑᡍٰՉ
१ނᒧǶԖࣴزஒԜҔٰࣴزނᢀჸᏢಞϐߏයᏫᐕำǴวεႵ ȐLong-Evans ratȑѝሡᆶҢጄޣอኩ 30 ϩដޑϕࡕ൩ૈԋჹҢጄޣමӞၸ ޑ१ނԖ܌ୃӳǴջ٬ҢጄޣςόӧǴ೭ኬޑᏢಞਏ݀ᗋࢂૈុঁД (Galef
& Whiskin, 2003)ǶӕኬճҔ೭ኬޑբǴӧаৎႵȐMus domesticusȑࣁނኳ
ࠠޑࣴز߾ࡰрǴԜᅿᢀჸᏢಞڙډޗ่ᄬޑፄᚇࡋԶቹៜޑǶҢጄޣ
ࢂᢀჸޣޑηж܈ޣࢂᆶᢀჸޣԃស࣬ӕޑќሟ܄Ǵ೭ᅿᏫёаӧҢጄޣς όӸӧࡕ 4 ঁλਔᗋૈߥ੮ǹՠӧ 24 λਔࡕǴѝԖᆶᢀჸޣԃស࣬ӕޑሟ܄Ң ጄޣ܌ԋޑᏫਏ݀ᗋӸӧǶӵ݀ஒҢጄޣޑኧໆቚӭǴॊٿᅿόӕҢጄޣ
܌ԋޑᏫਏ݀ǴёаӧҢጄޣςόӸӧࡕ 4 ঁλਔᗋૈߥ੮ǹՠࣣܭ 24 λ ਔࡕ࣮όډᏫӸӧޑ߄ (Choleris, Guo, Liu, Mainardi, & Valsecchi, 1997)Ƕ ΒǵᏢಞব္ёаפډ१ނ
ଞჹ೭ᜪޑࣴزᚒǴӭъࢂࣴز٩ᒘຎࣁЬޑނǶӧԜᚒ٠ؒԖ
ঁۓޑࣴزኳԄǴՠ୷ҁޑᏹБԄջࣁǺᡣᢀჸޣѐᢀჸҢጄޣӵՖפډ १ނǴฅࡕӆᡣᢀჸޣԾՉѐ൨פ१ނǶ೯தᢀჸޣᒧҔд܌ᢀჸၸޑБԄ
܈ᒧҢጄޣ܌ᒧޑӦᗺٰפ൨१ނǶ () аങᜪࣁኳԄ
ၸങᜪޑࣴزǴόѝวдॺёаၸᢀჸձޑҢጄޣޑ߄ٰפډ१ނ ޑ܌ӧǴᗋวፏӭቹៜᢀჸᏢಞޑӢનǶࣴزࡰрǴђКങȐPoecilia reticulataȑ
ୃӳၟᒿԖᡍǵዕޑǵឦܭεიᡏޑҢጄޣ (Lachlan, Crooks, & Laland, 1998)ǶᗨฅᢀჸޣୃӳၟᒿԖᡍޑҢጄޣǴՠό߄ҢૈԋؼӳޑᏢಞਏ݀Ƕ
ࣴزࡰрǴᢀჸޣၟᒿ߄ؼӳޑҢጄޣǴѝӧ໒ۈ߄Кၟᒿ߄ৡࠂޑ ҢጄޣٰளӳǹՠډΑࡕය໘ࢤǴϸԶᡂԋၟᒿ߄ৡޑҢጄޣޑᢀჸޣӧᏢ ಞೲࡋКၨז (Swaney, Kendal, Capon, Brown, & Laland, 2001)Ƕр೭ኬޑ
ຝǴࣴزޣᇡࣁǴࡐёૈࢂӢࣁ߄ؼӳޑҢጄޣϐՉǴߦ٬Αᢀჸޣஒݙ ཀΚܫӧቹៜ߄ޑᜢᗖՏ܈ڈᐟނǴԶ٬ளᢀჸޣӧ߃ۈ໘ࢤૈ߄ၨ
ӳǹ೭ᅿቹៜݙཀΚޑਏ݀Ξᆀࣁୱ܄மϯȐlocal enhancementȑ܈ڈᐟނம
ϯȐstimulus enhancementȑǶ࣬ჹٰᇥǴၟᒿ߄ৡࠂҢጄޣǴᗨฅ໒ۈݙཀ ΚޑᘍܫؒԖКၨזܫډ҅ዴՏǴՠҗܭόؼҢጄޣբКၨᄌǴᡣᢀჸޣ
ܰၟᒿǴԶЪԖ׳ӭᐒёаϩԖਏՉࣁᆶคਏՉࣁǴӢԜԋࡕය໘ࢤޑ ؼӳ߄ǹ೭ᅿਏ݀Ξᆀࣁёϩ܄ޑၟᒿՉࣁȐdiscriminated following behaviorȑǶќѦǴᗨฅᢀჸޣ഻ၟᒿεიᡏǴԶ߄рӵӕΓᜪШࣚύதـޑ வȐconformityȑǴՠᢀჸޣคݤӧຎ࣮ډԾρޑიᡏ܌ӧਔǴيೀܭε იᡏޑᢀჸޣᏢಞ߄߾Кيೀܭλიᡏޑᢀჸޣٰளৡǹᗨฅӆࡋܭຎૈ
࣮ډԾρޑიᡏ܌ӧਔǴيೀܭεიᡏޑᢀჸޣϐ߄ΞКيೀܭλიᡏޑᢀ ჸޣٰளӳ (Day, MacDonald, Brown, Laland, & Reader, 2001)Ƕ೭ኬޑຝΨས ҢΑவᗨฅԖշܭނှ،ढ़ғୢᚒǴՠࡐёૈቹៜཥڻՉࣁޑౢғǶவ
ॊϐࣴزᇥܴΑᢀჸᏢಞჹܭނӧפ൨१ނࢂख़ाЪڀቹៜ܄ޑǴόၸΨ ԖࣴزࡰрǴᢀჸޣ٠ߚښޑ٬ҔΓᡍǶӵ݀ঁᡏޑᡍ٠όሡाбрϼ εжሽਔǴКଆ٬ҔΓᡍǴᗋࢂ٩ᒘঁᡏᡍǹՠ٬Ҕঁᡏᡍሡбр εжሽਔǴ߾ᒧ٩ᒘΓᡍ (Kendal, Coolen, & Laland, 2004)Ƕӧΐڈങ ȐPungitius pungitiusȑޑࣴزΨวǴᢀჸޣᇡࣁ٬ҔԾيᡍǴԋфᕇள१ ނޑёૈ܄ၨଯਔǴКଆ٬ҔΓᡍǴᗋࢂ٩ᒘঁᡏᡍǹᢀჸޣᇡࣁ٬
ҔԾيᡍǴԋфᕇள१ނޑёૈ܄ၨեਔǴ߾٩ᒘΓᡍǶନԜϐѦǴ
ঁᡏޑᏫࢂຫΦϐޑᡍǴΨफ़եᢀჸޣ٬ҔԾيᡍޑёૈ܄ǴԶׯࣁ ٩ᒘΓᡍ (van Bergen, Coolen, & Laland, 2004)Ƕ
(Β) аܲᙝࣁኳԄ
ၸᢀჸᏢಞԶӧ१ՉࣁளǴၸѐᇡࣁᢀჸޣᆶҢጄޣࣁӕဂᡏǴ ཀջၨዕޑӕՔǴځਏ݀ၨӳǶόၸԖࣴزࡰрǴεီȐBombus impatiens CRESSONȑӧᢀჸΑձޑணҤޑҢጄޣϐ߄ǴΨрᆶҢጄޣኬޑᒧ
(Worden & Papaj, 2005)Ƕ೭ኬޑ่݀ᇥܴΑǴόӕޑဂᡏΨૈၸᢀჸᏢಞٰᕇ ளӳೀǴӆԛᡍᢀჸᏢಞڀഢΑᄽϯޑሽॶǶόၸ၀ࣴزΨᢀჸډǴ೭ኬޑ ਏ݀ջ٬ࢂҗΓπޑኳᔕҢጄޣΨૈၲډ࣬ӕޑਏ݀Ǵ՟ЯᇥܴΑ೭ኬޑՉࣁ߄
ǴёૈѝሡाൂપޑݙཀΚᐕำวғջёֹԋǶ
ມǵᆶ܄ᆶᕷ़ԖᜢޑᢀჸᏢಞՉࣁ
ଞჹ೭ᜪޑࣴزᚒǴӭъࢂࣴز٩ᒘຎޑނࣁЬǶ೭٤ࣴز೯த٬Ҕ
ଽ܈ҬଛϕኳԄǶ၀ኳԄޑ୷ҁБݤջࣁǺᡣᢀჸޣѐᢀჸሟวғ
ଽ܈Ҭଛϕ٣ҹǴϐࡕӆᡣᢀჸޣѐჹٿଫόӕܭᢀჸޣ܄ձޑڈᐟނᒧ
ǴځύଫؒԖᢀჸޣᢀჸၸԖวғၸଽ܈Ҭଛϕ٣ҹǴќଫ߾ࣁ
ᢀჸޣᢀჸၸԖวғ࣬ӕ٣ҹޑҢጄޣǶӧԜᚒΠ܌วғޑᢀჸᏢಞǴςόӆ
ࢂᢀჸޣᢀჸൂҢጄޣՉࢌՉԶவύᏢಞډҢጄޣޑՉǴԶࢂٿঁҢ ጄޣޑՉᢀჸᆶᏢಞǴӢԜ೭ኬޑࣴزǴჹܭዴᇡᢀჸޣ܌߄рٰޑՉࣁ
ࢂցᏢಞаϷவব္Ꮲಞډ၀ՉࣁࢂሡाߚதᙣޑǶ
ǵаങᜪࣁኳԄ
ӧђКങޑࣴزύࡰрǴሟ܄ᢀჸޣୃӳමᢀჸၸԖᆶځѬሟ܄Չၸ
ଽϕ٣ҹޑ܄ങǴЪᢀჸޣࢂᇡள၀܄ωՉԜᒧޑ (Dugatkin, 1992)ǶќѦǴᢀჸᏢಞޑቹៜΚόѝૈቹៜሟ܄ჹ܄ޑᄊࡋǴΨૈׯᡂচҁ ჹ܄ޑᄊࡋǶࣴزวԁ៧吃ങȐPoecilia latipinnaȑӧᢀჸډচҁ܌ୃӳ ޑ܄ځѬሟ܄ܔ㶥ǴΨӧϐࡕܔ㶥၀܄ǴԶ೭ኬޑᄊࡋׯᡂߚᒿᐒวғ ޑ (Witte & Ueding, 2003)Ƕ׳ᡋΓЪԖ፪ޑࢂǴᢀჸᏢಞ܌ԋޑਏ݀ջ٬ӧό ӕࠔᅿيΨૈวǶࣴزࡰрǴሟ܄ԁ៧吃ങୃӳӃᢀჸډԖჹќ
ᅿ௦ค܄ғБԄЪࣁόӕࠔᅿޑሟ܄٥ଭᇽԁങȐPoecilia FormosaȑՉ
ଽՉࣁޑ܄ԁ៧吃ങ (Schlupp, Marler, & Ryan, 1994)Ƕ ΒǵаചᜪࣁኳԄ
() ሟ܄ᢀჸޣ
ၸВҁȐCoturnix japonicaȑ܌ளډ่݀ᆶӧങᜪي܌ளډޑࣴز
่݀࣬ᜪ՟ (Galef & White, 1998)ǴନΑሟ܄ᢀჸޣѝሡᢀჸډ܄ᆶќଫ ሟ܄ઠӧଆ൩ૈჹ၀܄ౢғୃӳ (White & Galef, 1999a)Ǵ೭ኬޑ่݀Ψᇥܴ
ΑᢀჸډȸۓՉȹޑวғჹሟ܄ᢀჸޣനࡕޑᒧ٠ߚࢂѸाޑచҹǶќ ѦǴሟ܄ჹܭ܄ޑୃӳᗨฅ٬ளୃӳޣ٬ځڙѫޑᐒၨଯǴՠᢀჸᏢಞ ޑਏ݀ૈቚуόୃӳޣ٬ሟ܄ڙѫޑᐒගଯ (Persaud & Galef, 2005)ǶКၨ
Ԗ፪ޑࢂǴӵ݀܄ي܌ڀഢޑቻᆶሟ܄ᢀჸޣӃᢀჸډќଫමᆶځѬ ሟ܄Ҭଛၸޑ܄يޑቻ࣬ӕǴ߾ሟ܄ᢀჸޣΨჹڀԖ࣬ӕቻޑ܄Ԗ
܌ୃӳǴջ٬Ӄؒـၸ၀܄ (White & Galef, 2000)Ƕ (Β) ܄ᢀჸޣ
ӧᢀჸޣࣁ܄ਔ߾ளډᆶॊሟ܄่݀࣬ϸޑჴᡍ่݀Ǵ܄ϸԶ௨Ѿම ᢀჸډձޑ܄ଽၸޑሟ܄Ƕ೭ኬޑਏ݀ሡԖᢀჸډଽՉࣁޑрωӸ ӧǴΨᇥܴΑᢀჸډȸۓՉȹޑวғჹ܄ᢀჸޣനࡕޑᒧࢂѸाޑచҹ (White & Galef, 1999b)Ƕ
ୖǵᆶٛᑇԖᜢޑᢀჸᏢಞՉࣁ
ଞჹ೭ᜪޑࣴزᚒǴЬाӧࣴزঁᡏӵՖၸᢀჸᏢಞٰಞளૈᗉխӒᓀ วғޑՉǶ೭ᅿᚒޑࣴزБݤؒԖঁڰۓኳԄǴ೯தࢂᡣᢀჸޣѐᢀჸҢ ጄޣӵՖᚆঁܴᡉڀ࠶ુ܄ޑڈᐟނ܈ᕉნǶᗨฅૈᚆ၀࠶ુޑБԄёૈ
ԖࡐӭᅿǴՠᙖҗҢጄޣޑҢጄǴᢀჸޣஒёזೲᏢಞډࢌᅿᚆБԄǶ
ǵаങᜪࣁኳԄ
ӧђКങޑࣴزύวǴҢጄޣӧਔǴᢀჸޣ٬ҔҗҢጄޣٗᜐᏢ ಞٰޑၡጕٰזೲᚆ࠶ુ܄ڈᐟǹՠҢጄޣόӧਔǴᢀჸޣ܌٬Ҕϐᚆၡ ጕࠅёૈόࡪྣҢጄޣ߃ҢጄޑٗኬѐՉǴϸԶᒧΑќᅿૈ׳זೲ
ᚆ࠶ુ܄ڈᐟޑၡጕ (Brown & Laland, 2002)Ƕ೭ኬޑ่݀όѝᇥܴᢀჸޣᙖҗ
ᢀჸᏢಞளډӳೀǴΨӆԛསҢΑவቹៜཥڻՉࣁޑౢғǶ ΒǵаܲᙝࣁኳԄ
ӧᐋᖧȐNemobius sylvestrisȑޑࣴزύวǴᐋᖧӧᢀჸԖᡍޑӕᜪ߄
рӵՖၧᗉഁᇐȐPardosa spp.ȑޑՉࣁࡕǴᢀჸޣΨճҔӕኬБԄٰၧᗉ
ഁᇐޑрǴջ٬ҢጄޣςόӆрǴ೭ኬޑᏢಞ߄ᗋࢂёаុ 24 λਔ (Coolen, Dangles, & Casas, 2005)Ƕ೭ኬޑ่݀ᇥܴΑջ٬Ⴝᐋᖧ೭ၸߚဂۚ
ғࢲޑғނΨૈճҔᢀჸᏢಞٰᕇளӳೀǶ Οǵаথ٢ᜪࣁኳԄ
() а㖺ᏁᜪࣁኳԄ
ࣴزࡰрǴλႵȐCF-1ȑӧᢀჸӕᜪڀװᔐ܄ޑ૨ηѕࠈਔрੵภό
௵གϯȐanalgesiaȑޑຝǹԶᢀჸޣӧ 24 λਔࡕၶډ၀ۓᜪࠠޑ૨ηਔǴ ջ٬၀૨ηςܘନ֎ᆅԶคݤԋ࠶ુǴΨӵӕҢጄޣрஒيᡏশ
პޑၧᗉБԄǴՠᢀჸޣ܌߄рޑ೭ᅿՉࣁ٠όӧᢀჸᏢಞ໘ࢤύрӧ ᢀჸޣيǶନԜϐѦǴᗋрڋऊ܄ੵภό௵གϯȐconditioned analgesiaȑ ޑຝ (Kavaliers, Choleris, & Colwell, 2001)ǶࣴزᗋวǴၸ೭ኬޑȸᢀ ჸȹǴԋᢀჸޣޑҜ፦✉ȐcorticosteroneȑϲǹԶᢀჸޣӧ 24 λਔࡕၶ ډ၀ۓᜪࠠޑ૨ηਔǴջ٬၀૨ηςܘନ֎ᆅԶคݤԋ࠶ુǴΨౢғڋ ऊ܄Ҝ፦✉ϲޑຝ (Kavaliers, Colwell, & Choleris, 2003)Ƕ೭٤่݀ᇥܴ
ΑǴᢀჸޣޑዴӧჹวғӧӕᜪيޑ٣ҹՉᢀჸǶ (Β) аᒗࣁኳԄ
ࣴزύाᢀჸޣǴӧόޔௗаަᡏௗډҢጄޣޑΠǴᢀჸҢጄޣޑ ߄ࡕૈᏢಞډ᠋ډᖂॣૻဦ൩ाၢډጃηޑќᜐٰၧᗉႝᔐޑрǶ่݀
ᡉҢǴᢀჸޣಞளᗉՉࣁޑೲࡋזܭؒԖՉၸᢀჸᏢಞޑᒃيᐕޣǶԖ፪ ޑࢂǴᢀჸޣӧՉᢀჸᏢಞ૽ግǴමӧႝᔐጃύႝᔐၸԛ (John, Chesler, Bartlett, & Victor, 1968)ǴӢԜࡐёૈӃޑॄӛᡍǴ٬ளᢀჸޣჹܭ วғӧӕՔيӕኬޑᎁၶ׳௵ག܈׳ݙཀǴԶ٬ளᢀჸᏢಞޑਏ݀уமǶ
(Ο) аФࣁኳԄ
ԖࣴزࡰрǴϏഝើФȐDanish Friesian cowȑ߈ྕࢋޑྣ៝ޣǹჹ ಉኪޑྣ៝ޣ߾߄рᇻᚆޑՉࣁǶόၸᢀჸޣၸᢀჸӕՔаόӕБԄჹࡑ ٠คݤӢԜԶᏢಞډόӕྣ៝ޣϐྣ៝БԄǴᗨฅӧय़ჹӕՔࢂаྕࢋޑБԄ ჹࡑਔǴᒿӕՔᆶྣ៝ޣޑຯᚆຫ߈ǴΨ߄рຯᚆᅌ߈ޑՉࣁǶ೭ኬޑ่
݀ᗨฅؒԖࡰрФૈՉԜᅿࠠԄޑᢀჸᏢಞǴՠځՉࣁڙډӕՔޑՉࣁቹៜ (Munksgaard, DePassill, Rushen, Herskin, & Kristensen, 2001)Ƕ
(Ѥ) аญηࣁኳԄ
ӧԃᇸޑࡡݞญȐmacaca mulattaȑޑࣴزύวǴᢀჸޣၸӧቹТύᢀ ჸډӕᜪჹೂ߄р্܂ޑϸᔈࡕǴ߾চҁ߄рόឨ܂ೂޑԃᇸࡡݞญǴΨ
ӧϐࡕय़ჹೂޑрਔ߄р্܂ޑϸᔈǹՠӧӕኬޑᏹΠǴѝࢂஒቹТύޑ
ೂඤԋԚǴࠅόԋԃᇸࡡݞญѐჹԚр্܂ϸᔈ (Cook & Mineka, 1990)Ƕ೭ኬޑᏢಞᐒڋ܈ёаҔᢀჸڋऊȐobservational conditioningȑٰှញǴ ཀջᢀჸޣڙӕᜪޑ৮ឨᆣ܌གࢉǴԶΞௗय़ჹډচҁຎࣁύ܄ڈᐟނޑ
ೂǴӢԶౢғΑᙁൂᖄ่ᏢಞǶόၸൂൂаᙁൂޑᖄ่ᏢಞٰှញёૈΞόֹ
ӄǴӢࣁԃᇸࡡݞญ٠όᏢಞډा্܂Ԛ೭ᜪค্܄ޑނࠔǶ
စǵኳү
೭ᜪᚒޑࣴزाނѐՉسӈޑՉำׇǴӭъࢂࣴز٩ᒘຎ
ޑނࣁЬǶࣴزҞޑࣁၸᢀჸᏢಞǴёа٬ளᢀჸޣӧՉঁཥޑՉׇ
ӈਔ׳ࣁזೲ܈ҍၨϿޑᒱᇤǴᙖԜٰࣴزӸӧܭނيޑኳүȐimitationȑǶ ೯தჴᡍύ๏ϒނԋфՉՉޑዛ፞ࢂ१ނǴԶჴᡍޣᡣᢀჸޣೀܭঁ
ବᎧޑރᄊǴځҞޑӧቚуᢀჸޣޑᏢಞᐒᆶ߄ᐒǶZentall ӧ 2006 ԃޑ ӣ៝Ўകύᇡࣁा࠹ᆀނޑ߄ࣁኳүਔǴሡाૈڋȨޗߦȩȐsocial
facilitationȑǴཀջᢀჸޣᆶҢጄޣೀӧӕᕉნ܌ᏤठޑࢲΚϲǴԶࢲΚ ޑϲёૈԋՉଽฅֹԋǶᗋाૈڋӃ܌ගၸޑȨୱ܄மϯȩǵȨڈ ᐟނமϯȩǵᆶȨᢀჸڋऊȩǴаϷȨᇨวᏢಞȩȐaffordance learningȑǴཀջᢀჸ ޣၸҢጄޣޑՉԶᏢಞډڈᐟނ܈ᕉნޑфૈǶനࡕǴाዴᇡ೭٤Չಔԋ ӧচҁނيࢂόӸӧҭ܈ӧؒԖҢጄޣޑՉҢጄΠǴᢀჸޣԾฅр၀Չ
ޑёૈ܄ࢂࡐեޑȐZentall, 2006ȑǶ೭٤ӢનޑڋǴ٠ߚࡰ҅ޑኳүό
Ԗ೭٤ӢનӸӧǴԶࢂࡰ࠹ᆀࢌՉࣁ҅ޑኳүਔǴѸा௨ନҔ೭٤Ӣનջ
ૈֹӄှញ၀ՉрޑёૈǶӧࣴزނޑኳүǴؼӳޑڋಔीБԄЬा
ёϩٿᅿǺᅿࢂᅰёૈஒॊ܌ගӢન௨ନǹќᅿࢂȨᚈӛޑڋำׇȩ Ȑbidirectional control procedureȑǴ၀բाނӧࢌނᡏՉٿᅿόӕϸ ᔈǴԶόӕಔޑᢀჸޣஒ࣮ډҢጄޣՉόӕޑϸᔈǴӵԜ٬ள۶ԜջࣁჹБޑ
ڋಔǶ
ǵஒёૈӢન௨ନ
() аᒗࣁኳԄ
ӧᒗޑࣴزύวǴाᢀჸޣӧόޔௗаަᡏௗډҢጄޣޑΠǴᢀ ჸډҢጄޣޑ߄ࡕૈᏢಞډ᠋ډૻဦջаဌඓѐᓸఎηٰᕇள१ނǶ೭ኬ ޑՉाޑձϐೀӧܭނѝૈаȸဌඓȹѐᓸఎηǴӢԜࢂаۓيᡏ
ՏֹٰԋՉޑǶࣁΑዴᇡᢀჸޣޑԖᏢಞډ၀ՉǴ٠Ъ೭ኬޑᏢಞࢂፄ ᚇޑᐒڋǴӢԜࣴزύीΑٿঁڋಔǴځࢂᢀჸޣᢀჸډҢጄޣѝࢂ߈ ሺᏔ൩Ԗ१ނӞǴ೭ኬޑीࢂ௨ନݙཀΚᐕำวғ܌ᏤठՉࣁౢғޑёૈ܄Ǵ Ψ൩ࢂڋୱ܄܈ڈᐟނமϯǹځΒࢂᢀჸޣᢀჸډҢጄޣόᆅҔҺՖБԄᓸ ఎηԖ१ނӞǴ೭ኬޑीࢂӧዴᇡ၀Ꮲಞёૈόѝࢂᙁൂޑᖄ่ᏢಞǴКӵ ᢀჸڋऊǶ่݀ᡉҢǴԖᢀჸډҢጄޣҔဌඓѐᓸఎηޑКќѦٿಔڋಔ ߄р׳ӭ҅ዴᆶ׳זೲޑϸᔈ (John et al., 1968)Ƕ
(Β) а㖺ᏁᜪࣁኳԄ
ӧλႵ
Ȑ
C57BL/6Jȑ
ޑࣴزύǴाᢀჸޣӧόޔௗаަᡏௗډҢጄޣ ޑΠǴᢀჸҢጄޣޑ߄ࡕૈᏢಞډӃډሺᏔᜐᓸΠߎឦעЋ٬ᐒᜢ௴Ǵ१ނӢԜԶပډሺᏔޑΠቫܜஔࡕǴाາډሺᏔޑ҅БஒܜஔѺ໒ٰ
ڗள१ނ೭سӈፄᚇޑՉǴ೭ኬޑፄᚇՉࢂҗวғӧٿঁόӕՏޑٿ
ঁόӕՉࡪྣۓׇӈᜢ߯܌ಔԋޑǶ၀ࣴزӧᢀჸᏢಞ໘ࢤᆶᢀჸޣޑՉࣁ
ෳ၂໔ӼකΑঁਔ໔ޑۯ᪓Ǵ೭ኬޑीࢂࣁΑڋޗߦǴёа௨ନอኩ ࢲΚϲԶԋՉࣁрޑёૈ܄Ƕ่݀ᡉҢᢀჸޣԖᢀჸډҢጄޣޑՉ
ǴКଆคݤᢀჸډҢጄޣޑλႵǴૈ೯ၸ೭ፄᚇբޑКٯၨଯ (Carlier
& Jamon, 2006)Ƕ Βǵᚈӛޑڋำׇ
() а㖺ᏁᜪࣁኳԄ
ӧаεႵ
Ȑ
hooded Lister ratȑ
ࣁኳԄޑࣴزύǴ૽ግҢጄޣஒਥఎη ۳ѓ܈۳ѰǴԖъޑᢀჸޣஒ࣮ډҢጄޣעఎηӛѰǴќъ߾࣮ډ ఎηӛѓǶ่݀ᡉҢᢀჸޣ߄рдॺ܌ᢀჸډޑՉǶࣁΑ௨ନԴႵࢂճҔఎηර܊໔ύόӕय़ӛᕅᏛٰ౽ࣁጕǴӢԜࣴزޣӧᢀჸޣޑᢀჸ ໘ࢤ่״ࡕஒఎηޑᘍܫՏ౽ډќय़ᕅᏛǴ٬ளఎηޑ౽Бӛவচҁޑ ѰѓᡂԋѝૈࡕǶ่݀ӧෳ၂໘ࢤਔᢀჸޣᗋࢂ߄р܌ᢀჸډޑՉ
(Heyes & Dawson, 1990)Ƕ (Β) аചᜪࣁኳԄ
а㖺ᏁᜪࣁኳԄౢғ٤ୢᚒǴջคݤ௨ନԴႵޑ߄ёૈѝࢂᅿȨᇨ วᏢಞȩ܈ޣԴႵёૈճҔූ੮ܭఎηޑښၰጕǶࣁΑှ،ӵԜୢᚒǴ
ࣴزޣճҔӕኬޑБԄࡼෳӧ٩ᒘຎޑނيǴӵВҁǴӢԜ൩όԖ ښၰූ੮ޑୢᚒǶᢀჸޣஒӧᢀჸᏢಞය໔ᢀჸډߐҢጄޣරόӕБӛ౽Ƕ ჴᡍύቚуќڋಔǴᢀჸޣӕኬᢀჸډߐරόӕБӛ౽ǴѝࢂԜ౽όࢂ
Ңጄޣ܌ᏤठޑǶ่݀ᡉҢǴᢀჸޣ٩܌ᢀჸډߐޑ౽БԄѐՉǹՠӧ
ڋಔ߾όǶӢԜȨᇨวᏢಞȩҭคݤֹӄှញᢀჸޣ܌߄рٰޑՉࣁ (Akins, Klein, & Zentall, 2002)Ƕ
ಃΟʳ ޗ܄ૻ৲᠍ڗ
ኳүΨࢂᢀჸᏢಞޑᅿǴόၸኳүமፓޑࢂᢀჸޣၸᢀჸԶಞளҢጄޣ ޑȸՉȹǴՠᢀჸޣځჴёаѝࢂၸᢀჸҢጄޣޑՉԶಞளȸૻ৲ȹǶၸ ѐჹނޑᢀჸᏢಞՉࣁࣴزǴӭࢂӧނᙖҗᢀჸᏢಞޑБԄٰᕇளࢌᅿ ՉǴՠӵ݀ाჹᢀჸᏢಞ೭ኬޑᐕำԖ׳ӭޑΑှǴךॺόૈ۹ౣќᅿಞள ޑϣǺૻ৲Ƕाࣴز೭ᚒǴ܈ၸࣴزނԾฅวғޑޗ܄ᝡݾՉࣁǴ ёаᡣךॺჹ೭ᅿȸૻ৲ȹಞளޑᢀჸᏢಞԖ׳ӭޑΑှǶ
൘ǵวғܭޗ܄ᝡݾύޑޔௗຑᆶ໔ௗຑ
ӧғނ໔Ǵӕᜪ໔۶ԜϕᝡݾࢂࡐีѳதޑǴځҞޑεӭࢂࣁΑݾၗ
ྍǴ೭٤ၗྍёૈࢂ१ނޑٰྍǵҬଛޑΚǵ܈ޣ৲ӦޑጄൎǶՠӵࣁΑݾ
ၗྍǴԛЇଆᐟਗ਼ޑѺରǴჹނٰᇥࢂڀ॥ᓀ܄Ϸ্܄ޑǴӢࣁᐟਗ਼ޑ ѺରёૈᏤठڙࣗԿܭࢂԝΫǶ܌аނӧᝡݾύሡჹӕᜪޑᏯରૈΚՉຑ
ǴѺରၸำύຑрჹЋૈΚКԾρமၸӭǴ൩όा੮ᠰܭၗྍݾڗǴᔈ
ᅰזܫకᏯରќ൨рၡǶՠঃऩሡाၸԛჴሞᏯରᐕωૈຑჹЋޑૈ
ΚǴԶคݤڗ௳ၸޑ௲૽Ǵ߾ڙޑ॥ᓀᗋࢂӸӧǶӢԜނाૈᏢಞ ډӧۓᡍΠՉঁᡏӆᇡȐindividual recognitionȑǴཀջૈϩჹЋࢂցම
௳ၸԾρǴӵ݀ӆᇡрჹЋම௳ၸԾρǴ߾ᔈᒧόᏯԶଏǶၸѐӧ
௶ճ٥ঊႵȐMesocricetus auratusȑޑࣴزύ൩ࡰрǴ܄ঊႵӧ௳ࡕǴӵ
ӆࡋޔௗय़ჹ࣬ӕޑჹЋਔόᏯԶǹନԜϐѦǴჹ၀ۓჹЋޑᏫёа
ុԿϿΎϺ (Lai & Johnston, 2002)Ƕՠ٣ჴǴջ٬೭ኬૈᏢډ௲૽Ǵёૈᗋࢂ
ڀԖۓޑ॥ᓀ܄ǴӢࣁሡाᒃيᐕᏯରωૈֹԋຑǶ٣ჴǴӵӕӃჹ วғܭނيޑᢀჸᏢಞՉࣁ܌Չޑӣ៝ǴᢀჸᏢಞࢂڀԖᄽϯϷނғ Ӹख़ाޑሽॶǴӢԜךॺёаෳǴନΑၸޔௗޑᒃيᐕٰՉእЋຑ
ѦǴᔈ၀ӸӧૈၸᢀჸдΓޑᡍٰᕇளૻ৲ǴԶၲډᕭอእЋຑᐕำ ޑБԄǶ٣ჴᡉҢǴၸഁӦᢀჸᆶჴᡍ࠻ࣴزǴ೭ᅿၸΓޑᡍٰၲډૻ
৲ᕇளޑБԄࢂӸӧޑǴᆀࣁȨޗ܄ૻ৲᠍ڗȩȐeavesdroppingȑǶ
ޗ܄ૻ৲᠍ڗຒ٬ҔٰඔॊނၸᢀჸќѦٿଫӕᜪޑϕԶவ ϕύௗԏډૻ৲ޑᅿБԄ (McGregor, 1993)ǶEavesdrop ೭ঁຒࢂҗ eavesdripᙯᡂԶٰǶEaves ࢂࡂᛞޑཀࡘǹԶ drip ࡰளࢂᅀΠǴӢԜ eavesdrip ӝଆٰࡰޑ൩ࢂࡂᛞஒߘНௗՐޑ೭ᅿБԄǴԶӵԜБԄԋߘНࡂᛞޑ
ྎࢬډࡂѦѐǶҗܭђжהޣୂ᠋ਔஒԾρॹӧࡂᛞǴ൩ӳႽߘНࢬΠޑኬ ηǴࡕٰ೭ᅿୂ᠋ޑՉࣁΨ൩ᆀࣁ EavesdroqǴԶ೭ኬޑཷۺ൩ۯ՜ډ೭ᅿ
ࣴزૻ৲ᕇளޑᚒύǶޗ܄ૻ৲᠍ڗޑ၁ಒۓကࣁǴၸᢀჸќѦٿଫӕᜪ ޑϕǴԶவϕޑૻဦύѐڗрૻ৲٠ӧ҂ٰѐ٬Ҕϐ (McGregor, 1993;
McGregor & Peake, 2000)Ƕӧϕύ܌х֖ޑૻ৲ȐinformationȑǴКБӵᏯରૈ
ΚǴࠔ፦ǴᐒǴ೭٤ૻ৲όࢂҗૻဦȐsignalȑҁي܌ಔԋǴԶࢂӸӧܭ ೭٤ૻဦ࡛ሶ٬Ҕӧ೭ϕϐύǶ೯தाബঁޗ܄ૻ৲᠍ڗޑᕉნǴന ᙁൂޑБԄ൩ࢂᡣӕ܄ձ܈ޣόӕ܄ձޑᢀჸޣѐᢀჸќѦٿଫӕ܄ձޑނ
Չᝡݾ܄ϕ٣ҹǶाዴᇡᢀჸޣࢂցςᕇளૻ৲Ǵ߾ሡाѐෳໆᢀჸޣӧᢀ ჸ٣ҹวғϐࡕǴय़ჹ܌ᢀჸډޑٿঁҢጄޣޑϸᔈࢂցԖ܌όӕǴӵ݀Ԗόӕ ջё࠹ᆀᢀჸޣςճҔޗ܄ૻ৲᠍ڗޑБԄᕇளૻ৲ǶӧԜᚒаނࣁኳ
ࠠޑࣴزςԖऩυЎ่݀ǴԖ٤ࣴز಄ӝޗ܄ૻ৲᠍ڗޑۓကǹՠԖ٤߾ό
಄ӝǴՠၸΑှ೭٤ࣴزύޑࣴزౣᆶ܌ளډޑ่݀ǴஒԖշܭךॺჹ೭
ᚒޑǶ
ມǵޗ܄ᝡݾύଞჹޗ܄ૻ৲᠍ڗӧނيޑࣴز
ǵഁӦࣴز
ԐයഁӦࣴزࡰрǴ܄आવ㚁ȐAgelaius phoeniceusȑӧيޑᎃۚ
ѦእߟΕԾৎሦӦਔǴߟΕޣࢂଫόڀװᔐ܄ޑܿऍࢪЦ៙㒣ȐTyrannus
tyrannusȑǴᢀჸޣߟҍ၀ᎃۚሦӦޑՉࣁ൩൳ЯؒวғǹՠӵߟΕޣࢂќଫڀ
װᔐ܄ޑ܄आવ㚁Ǵᎃۚӧय़ჹ೭ᅿѦእߟΕਔ܌๏ϒޑװᔐၨ১ਔǴ߾
ᢀჸޣߟҍԜۓᎃۚሦӦޑՉࣁ൩ၨதวғ (Freeman, 1987)Ƕ೭ኬޑ่݀
՟Я໔ௗᇥܴ܄आવ㚁ёૈၸȸᢀჸȹٰᏢಞᎃۚޑ០እૈΚǶ Βǵჴᡍ࠻ࣴز
() аങᜪࣁኳԄ 1. ܄ᢀჸޣ
ӧа܄ᐁᛥରങȐBetta splendensȑࣁኳԄޑࣴزύǴࣴزޣࢂᡣᢀჸޣ ѐᢀჸٿଫ܄ങӧϕϣՉ 5 ϩដޑᝡݾ܄ϕǶၸܴ႖݈ஒ۶Ԝ
႖໒ٰǴ٬ளᢀჸޣᆶٿଫҢጄޣӅΟଫ܄ങǴคݤӧަᡏՉޔௗௗǴ ՠૈၸຎᆶ༘ޑБԄٰՉૻဦᆶૻ৲ޑሀǶӧ೭ኬޑჴᡍᏹΠǴࣴ
زޣԖᢀჸډǴᝡݾ܄ϕՉϐਔǴᢀჸޣय़ჹԖ٣ҹวғޑٗঁБӛǴ ӢԜࣴزޣᇡࣁᢀჸޣԖӧჹ٣ҹՉᢀჸǶӧᢀჸᏢಞ໘ࢤ่״ࡕǴࣴزޣ
ஒϕύޑᒡৎȐloserȑڗрǴ੮ΠគৎȐwinnerȑᆶᢀჸޣǴ႖Ϻ߾ՉՉࣁ
ෳ၂໘ࢤǶՉࣁෳ၂ᕉნࢂќঁཥᕉნǴគৎᆶᢀჸޣᒿᐒϩԋٿಔǴ
ಔᢀჸޣᆶচٰ܌ᢀჸၸޑគৎҢጄޣଛჹǹќಔᢀჸޣ߾ᆶќଫؒـ ၸՠᡏख़ᆶځ࣬ৡόӭޑគৎҢጄޣଛჹǶঁଛჹܫӧϕϣǴӧؒԖ
ܴ႖݈ޑΠՉޔௗ܄ޑᝡݾ܄ϕǴࣴزޣ߾ஒϕރݩᒵΠٰǶࣴ
ز่݀ࡰрǴᢀჸޣόፕനࡕࢂയࢂॄǴӧ،ۓाᐟᏯ܈ޣाܫకǴӧᆶዕ
ޑគৎҢጄޣՉϕКଆᆶढ़ғޑគৎҢጄޣՉϕޑ೭ٿᅿნΠǴᢀჸ
ޣӧᆶޣޑϕϐՉ،׳זೲǴᡉҢӃޑᢀჸૈ෧ϿእЋૈΚຑ
ޑਔ໔Ǵٰᡣঁᡏ׳זೲрղᘐ (Johnsson & Ækerman, 1998)Ƕ೭ኬޑࣴز่
݀КଆഁӦᢀჸளٰޑ่݀Ǵ׳ԖᐒܴᢀჸޣԖӧჹᝡݾ܄ϕ٣ҹՉᢀ ჸǶόၸǴջ٬ӵԜǴ၀ࣴزᗋࢂӢࣁჴᡍीޑόֹഢԶคݤܴዴܴᢀჸ ޣԖӧჹᝡݾ܄ϕ٣ҹՉᢀჸǴӢࣁڙ၂ޣරӛԖ٣ҹวғޑٗঁБӛǴё
ૈѝࢂӢࣁٗঁБӛԖӕᜪޑӸӧǶӕԃޑќঁࣴزύǴ߾Αၨ٫ޑڋٰ
ዴᇡԜ٣ǶࣴزޣஒჴᡍሺᏔϩԋᢀჸᆶϕǴϕࢂӧᢀჸޑѰѓٿ ᜐǶᢀჸޣӼ௨ᢀჸځύᜐޑϕύԖٿଫҢጄޣၸܴ႖݈ӧՉ ໔ௗޑᝡݾ܄ϕ٣ҹǹќᜐ߾ᢀჸډٿଫҢጄޣӢࣁύ໔႖όܴ႖݈Զ
ؒԖӧՉϕǶҗܭҢጄޣᆶᢀჸޣ܌ೀޑୱԖܴߝࡋޑৡ౦ǴӢԜᢀჸ ޣૈᢀჸډҢጄޣޑࢲǴՠԾيόҢጄޣ๏ᢀჸډǶӧ೭ኬޑीΠǴ၀ࣴ
زࡰрǴ࣬ჹܭؒԖᝡݾ܄ϕ٣ҹวғޑᜐǴᢀჸޣჹܭԖวғᝡݾ܄ϕ
٣ҹޑٗᜐݙຎၨΦǴᡉҢΑᢀჸޣޑዴԖӧݙཀᝡݾ܄ϕ٣ҹޑวғǶ
ௗࣴزޣགྷዴᇡᢀჸޣࢂցவᝡݾ܄ϕ٣ҹޑᢀჸύಞளૻ৲ǴӢԜ௦ҔΑ ќᅿीǶᢀჸޣӼ௨ᢀჸځύᜐޑϕԖٿଫҢጄޣၸܴ႖݈
ӧՉ໔ௗޑᝡݾ܄ϕ٣ҹǹќᜐ߾Ӣόܴ႖݈ߔ႖ǴᏤठคݤᢀჸډ ќѦٿଫҢጄޣӕኬӧՉ໔ௗޑᝡݾ܄ϕ٣ҹǶᢀჸᏢಞ໘ࢤ่״ࡕଭ
ՉՉࣁෳ၂໘ࢤǴ܌ԖޑҢጄޣϩձးӧঁܴ౯ηύǴаԛଫޑ БԄΕᢀჸޣ܌ࡑޑୱϣǴࣴزޣ߾ෳໆᢀჸޣჹߟΕޣޑϸᔈǶӧ೭ኬޑ
ीΠǴ၀ࣴزࡰрǴ࣬ჹܭय़ჹᢀჸډᒡၸдΓޑΕߟޣǴᢀჸޣჹܭᢀჸ ډគၸдΓޑΕߟޣ߄рᄌޑᖿ߈Ǵ٠ЪၨᄌωჹځՉ࠶ᓵǹՠ೭ኬޑ ৡ౦܄ӧόᡣᢀჸޣԖᐒᢀჸډҢጄޣॺ܌Չޑᝡݾ܄ϕ٣ҹਔ൩ό
рǶ೭٤่݀ᡉҢΑڙ၂ޣޑዴԖӧჹӕᜪޑᝡݾ܄ϕ٣ҹՉᢀჸǴ٠வ ύಞளૻ৲ǴЪஒ٤ૻ৲ࣁࡕុᏯରӝޑ߃ۈՉ٩Ᏽ (Oliveira, McGregor,
& Latruffe, 1998)Ƕჹܭዴᇡᢀჸޣ܌ಞளૻ৲ࢂٰԾܭӕᜪޑᝡݾ܄ϕϐ ύǴନΑճҔނ܌วғޑჴᏯରǴᗋԖࣴز٬ҔΑќᅿѯ֮ޑीٰዴᇡ
Ԝ٣ǶࣴزύǴࣴزޣᡣᢀჸޣൂБय़ᢀჸډϕԖٿଫҢጄޣǴA ᆶ B ၸܴ႖݈ӧՉ໔ௗޑᝡݾ܄ϕ٣ҹǶѯ֮ޑࢂǴᝡݾ܄ϕ٣ҹύځჴࢂ
ٿٿ܄ങӧ۶ԜՉϕǴA ᆶ B аϷ C ᆶ DǶࣴزޣஒ B аϷ C ᗦᙒՐǴ
٬ளவᢀჸޣޑفࡋٰ࣮ǴӢࣁѝ࣮ډ A ᆶ DǴԶᇤаࣁࢂ A ᆶ D ӧՉϕǶ ӧᢀჸᏢಞ໘ࢤ่״ࡕǴࣴزޣϩձᡣᢀჸޣᆶ A ܈ B Չ໔ௗޑᝡݾ܄ϕ
ǴӕኬෳໆᢀჸޣჹჹЋޑϸᔈǶӧ೭ኬޑीΠǴࣴزࡰрǴ࣬ჹܭय़ჹᢀ ჸډȸᒡၸдΓȹޑჹЋǴᢀჸޣჹܭᢀჸډȸគၸдΓȹޑჹЋၨזޑᖿ߈ аϷၨזޑჹځՉ࠶ᓵǶҗܭӧ೭ኬޑीΠǴ٬ளჴШࣚ܌วғޑ٣ҹᆶ ᢀჸޣ܌ޕډޑ٣ҹࢂόӕޑǴӢԜᢀჸޣჹόӕჹЋ߄рόӕՉࣁǴѝૈ
ᘜӢܭ೭ٿঁჹЋӧᢀჸޣޑΠޑ࣬ჹ߄܌Ꮴठޑ (McGregor, Peake, &
Lampe, 2001)Ƕ
2. ሟ܄ᢀჸޣ
ޗ܄ૻ৲᠍ڗΨૈၮҔӧሟ܄ϐଛଽᒧǶࣴزวǴሟ܄ᐁᛥରങӧ य़ჹԖวғᝡݾ܄܈ؒวғᝡݾ܄ϕ٣ҹ߄рόӕՉࣁǴԶჹܭԖวғ ᝡݾ܄ϕ٣ҹޑٿঁ܄ڈᐟނǴᢀჸޣᒧӃ߈គৎҢጄޣǴ٠ၨӭ ਔ໔ݙຎаϷ߄рၨӭޑғՉࣁǴ՟ЯКଆᒡৎҢጄޣǴሟ܄ᢀჸޣКၨୃ
ӳគৎҢጄޣ (Doutrelant & McGregor, 2000)Ƕ (Β) аചᜪࣁኳԄ
1. ܄ᢀჸޣ
ࣁΑؼӳڋϕၸำаϷஒૻ৲ޑӸӧᆒዴڋӧϕύǴԖᅿჴᡍЋ ݤ൩ࢂճҔኞܫᏔѐኳᔕٿଫചᙖҗሳћٰՉᝡݾ܄ϕ٣ҹǴԶ٣ҹՉ ύǴԖќଫჴޑ܄ചӧՉᢀჸǶࣴزࡰр܄εξഒȐParus majorȑ όፕࢂჹᝡݾ܄ϕ٣ҹวғޑᢀჸΠ܈ޣय़ჹаኞܫᏔኳᔕԋޑΕߟޣޑ ߟౣǴ߄рזೲᖿ߈ޑՉࣁǴԶЪࡕޣКޣᗋٰள׳זǶჹܭΕߟޣޑ ߟౣǴԜߟΕޣӃᢀჸޣ᠋ډӧᝡݾ܄ϕ٣ҹύࢂឦܭۚܭ১༈ޑ
БǴ߾ჹѬԖၨϿޑӣᔈǴᡉҢᢀჸޣૈᙖҗᢀჸӕᜪ໔ޑ۶ԜሳћރᄊǴٰ
ಞளӕᜪ໔ޑՏ໘ᜢ߯ (Peake, Terry, McGregor, & Dabelsteen, 2001)Ƕ 2. ሟ܄ᢀჸޣ
ӧаሟ܄ᐁᛥରങࣁኳԄ܌ளډޑ่݀ǴӧаВҁࣁኳԄࠅؒԖளډ
࣬ӕ่݀ǶࣴزࡰрǴӧᢀჸၸᝡݾ܄٣ҹޑวғࡕǴሟ܄ᢀჸޣᒧۚΠ॥
ޑ܄ڈᐟނǶࣴزޣჹܭ೭่݀ޑှញࢂǴሟ܄ёૈाᗉխᏱԖଯװᔐ܄ޑ
܄ӧՉҬଛਔװᔐǹ၀ࣴزΨวǴຫڀᓬ༈ޑ܄ǴຫӧՉҬଛਔ
р্܄ޑբ (Ophir & Galef, 2003)Ƕ
ୖǵঁᡏᡍჹޗᝡݾύޑޗ܄ૻ৲᠍ڗޑቹៜ
ᢀჸޣӧՉእЋຑϐਔǴၸѐ܌ᏱԖޑΓᡍ՟Я՞Ԗख़ाޑቹៜ ΚǶόၸࣴزΨวǴᢀჸޣόѝёаճҔΓᡍǴᗋёаӝٳ٬Ҕঁᡏᡍ ᆶΓᡍ೭ٿᅿόӕٰྍޑૻ৲Ƕ၀ࣴزӕኬճҔኞܫᏔѐኳᔕٿଫ܄εξ ഒᙖҗሳћٰՉᝡݾ܄ϕ٣ҹǶჴᡍϩΟঁ໘ࢤǴӧಃ໘ࢤǴᢀჸޣȐSȑ
ௗڙঁᡏᡍޑᏹǴёૈࢂ௳ȐA > Sȑ܈ޣࢂᏯയߟΕޣȐA < Sȑǹӧ
ಃΒ໘ࢤǴᢀჸޣௗڙΓᡍޑᏹǴӢԜᢀჸډӃޑߟΕޣᆶќჹ ЋՉᝡݾ܄ϕȐA > B ܈ A < BȑǹӧಃΟ໘ࢤǴӧಃΒ໘ࢤ܌ᢀჸډޑќ
ჹЋȐBȑஒߟΕᢀჸޣޑሦӦǶ่݀ࡰрǴᢀჸޣၸٿᅿόӕૻ৲ٰྍޑ ӝٳԶஒߟΕޣղᘐࣁόڀ࠶ુਔȐS > A > BȑǴჹ၀ߟΕޣԖၨϿޑӣᔈǹ ՠӵղᘐࣁڀ࠶ુ܄ȐB > A > SȑǴ܈ޣၗૻόىคݤΠղᘐਔȐS > A < B ܈ S <
A > BȑǴ߾Ԗၨӭޑӣᔈ (Peake, Terry, McGregor, & Dabelsteen, 2002)Ƕ೭ኬ ޑຝࢂ࣬Ԗ፪ޑǴӢࣁ೭ኬޑ߄སҢޗ܄ૻ৲᠍ڗ೭ૈΚว චډཱུठਔёૈрȨሀ౽ȩȐtransitive inferenceȑǴӕਔᡉҢ೭ᅿຎࣁ ଯᇡޕޑ߄ΨૈၸᢀჸޣᏢಞޑБԄٰՉǶӧሦβങȐAstatotilapia burtoniȑޑࣴزύ൩วǴᢀჸޣӧೱុᢀჸჹΞჹޑଛჹᝡݾ܄ϕ٣ҹ
ࡕǴёૈૈрჹЋ AǵBǵCǵDǵаϷ E ϐ໔Ӹӧ A > B > C > D > E ޑՏ໘ᜢ߯ǶӢࣁջ٬ᢀჸޣؒᢀჸၸჹЋ A ᓬܭ E ܈ B ᓬܭ D ޑଛჹಔӝǴ Ψ߄рᇻᚆჹЋ A ՠᖿ߈ E ޑՉࣁ܈ޣᇻᚆ B ՠᖿ߈ D ޑՉࣁ (Grosenick, Clement, & Fernald, 2007)Ƕ
ಃѤʳ ҁፕЎޑࣴزᚒᆶౣ
ၸޗ܄ૻ৲᠍ڗ೭ᅿᢀჸᏢಞޑБԄٰಞளૻ৲Ǵ٬ளঁᡏӧ٬ҔΓ
ᡍਔǴӭΑᅿঁᡏԾՉᒧޑޜ໔Ǵঁᡏёаᒧा٬Ҕ܈ޣό٬ҔǴࣗԿ ӝٳځѬٰྍޑૻ৲ࡕӆ،ۓǴӢԜךॺᇡࣁ೭ኬޑᏢಞБԄࢂᅿଯᇡޕ
ૈΚޑ߄ǶќѦǴޗ܄ᝡݾӧғނ໔ࢂࡐதـޑϕ٣ҹǴࣴزวғӧޗ
܄ᝡݾϕύޑޗ܄ૻ৲᠍ڗࢂుڀғᄊਏࡋޑǶၸѐଞჹޗ܄ૻ৲᠍ڗ೭ ᅿࠠԄޑᢀჸᏢಞǴӭъࢂ٬Ҕങᜪ܈ࢂചᜪࣁނኳԄǴ௦ҔΓᜪаѦޑځ Ѭথ٢ᜪϐࣴز߾ࢂϿϐΞϿǶӢԜӵ݀ૈճҔ㖺Ꮑᜪ೭ኬ࣬ჹКၨଯЪௗ߈ ΓᜪޑނᅿǴࣗԿࢂ㖺ᏁᜪჴᡍނǴٰࡌҥрཥޑޗ܄ૻ৲᠍ڗϐނኳ ԄǴନΑቚךॺჹόӕނᅿӧԜૈΚޑΑှѦǴΨԖշܭךॺճҔԜჴᡍ
ނኳԄѐᆶᙶమԜՉࣁङࡕޑတઓᐒڋǶ
൘ǵࣴزᚒ
ǵޗ܄ૻ৲᠍ڗᆶ௶ճ٥ᅿঊႵ
௶ճ٥ঊႵ೭ᅿ㖺ᏁᜪނǴᇡࣁԋԃࡕӧഁӦύࢂՉᐱۚޑғࢲΔӧჴ ᡍ࠻ύԋԃࡕΨӭаᐱҥޜ໔ޑБԄஒ۶Ԝϩ໒ႩᎦǶԋԃঊႵჹܭځѬΕߟ ޣڀԖଯװᔐ܄ǴࣁΑٛፁԾρޑࢰҤᆶሦӦԶᆶځѬӕᜪวғᏯରǶόၸ ঊႵ٠ߚрғ൩ࢂᐱۚǴ೯தӧᚆணࡕ܈ࢂࢌঁ໘ࢤࡕǴω໒ۈځᐱۚғࢲǴ
ՠջ٬ՉᐱۚғࢲǴځࢲሦୱΨᆶځдӕᜪԖۓำࡋޑख़᠄ᆶϕǶԖࣴ
ز൩ࡰрǴᐱۚ܄ၨ১ޑԋԃࣔࠠঊႵȐPhodopus campbelliȑӧαښୃӳޑ ޗሀբޑᢀჸᏢಞਏ݀ၨӳǴՠ࣬ჹٰᖱᐱۚ܄ၨமޑԋԃ௶ճ٥ঊ Ⴕ߾ѝԖҢጄޣࢂᢀჸޣޑ҆ᒃǴ٠Ъᢀჸޣԃइऊ 4 ঁࢃයεਔωૈว
ԖᢀჸᏢಞޑਏ݀ǶόၸКଆόࢂவҢጄޣ္ڗрޑ१ނǴԋԃ௶ճ٥ঊႵ
ୃӳᒧவҢጄޣ္ڗрޑ१ނ (Lupfer, Frieman, & Coonfield, 2003)Ƕᇥܴ
ΑԃѴޑ௶ճ٥ঊႵڀԖۓޑᢀჸᏢಞૈΚǹԶԋԃޑঊႵǴӧᒧ१ނ
ᗋࢂёаၸᢀჸᏢಞٰಞளǴѝࢂӢࣁલЮᒃஏޗϕޑᐒ܈ࢂࣁΑᗉ խόѸाޑѺରǴӢԶफ़եΑ೭ᜪᢀჸᏢಞวғޑᐒǴՠϝฅڀഢۓำࡋ ޑᢀჸᏢಞૈΚǶҗܭԋԃঊႵڀԖሦୱ܄ՉࣁǴӢԜၶډΕߟϐঊႵǴ൩ ёૈวғѺରՉࣁǶ೭ᅿ܄ςԋфճҔٰᡍр܄ঊႵࢂڀԖঁᡏӆ ᇡޑૈΚǴΨ൩ࢂૈӆᇡрѺគၸԾρޑۓእΓ (Lai & Johnston, 2002)Ƕӧ ଛଽᒧϐᢀჸᏢಞࣴزΨࡰрᢀჸᏢಞੋϷΑঁᡏӆᇡޑᐕำ (Dugatkin, 1992)ǶӢԜճҔԋԃ܄ঊႵԖሦӦٛᑇՉࣁǵёаӧჴᡍ࠻ൂᐱႩᎦᗉխό ѸाυᘋǵуڀԖૈঁᡏӆᇡޑૈΚǵаϷڀԖۓำࡋޑޗϕᆶᢀჸ ᏢಞૈΚᓬᗺǴԖёૈԖᐒёаӧԋԃঊႵᢀჸډޗ܄ᝡݾϕύޑ ޗ܄ૻ৲᠍ڗϐᏢಞБԄǴ٠ࡌҥჴᡍᏹբϐՉࣁኳࠠǶ
Βǵঁᡏᡍᆶޗ܄ૻ৲᠍ڗ
ӧᒗޑᢀჸᏢಞࣴزύǴࣴزޣၸ٣Ӄ๏ϒᢀჸޣԛႝᔐᡍࡕω
ՉᗉႝᔐޑᢀჸᏢಞ૽ግǴԖวᢀჸᏢಞਏ݀ޑр (John et al., 1968)Ǵ՟
ЯӃޑॄӛᡍߦࡕុޑᏢಞਏ݀ǶόၸǴ၀ࣴز٠ؒԖᏹคӃঁᡏ
ᡍΠޑᢀჸᏢಞნǴӢԜόዴۓӃޑॄӛᡍࢂցቹៜࡕុޑᢀჸᏢಞ ᐕำǴՠၸа܄εξഒࣁኳࠠӧޗ܄ૻ৲᠍ڗᚒ܌ޑࣴزǴ໔ௗ࣮
ډᢀჸޣӃޑόӕঁᡏᡍǴቹៜࡕុځ٬ҔΓᡍޑБԄǴᗨฅ၀ࣴز ჹܭᢀჸޣӧΓૻ৲ᕇளϐਔԖՖϸᔈ٠คගϷ (Peake et al., 2002)ǶӢԜ
ፕǴӃޑঁᡏॄӛᡍǴٯӵޗҬ௳Ǵᔈ၀Ψჹวғܭޗ܄ᝡݾϕύ
ޑޗ܄ૻ৲᠍ڗϐᢀჸᏢಞᐕำԋቹៜǴࣗԿቹៜډࡕុޑᏢಞ߄Ƕ ΟǵᓸΚᅟᆾǵᓸΚೀᆶޗ܄ૻ৲᠍ڗ
Չޗ܄ᝡݾϕǴёૈԋނڙǴӢԜ೭ኬޑ٣ҹჹܭނٰᇥ ёૈ൩ࢂڀᓸΚޑǴԶيᡏӵՖჹ೭ኬޑᓸΚՉঁӝޑೀǴ൩ᡂԋࡐख़
ाЪࡐჴҔޑᚒǶΠຎЫ-တࠟᡏ-Ҝ፦ືጕȐhypothamamic-pituitary-adrenal cortex axisȑǴӧথ٢ᜪނय़ჹᓸΚნ܈ڈᐟޑೀᆶᔈתᄽख़ाف ՅǶϩݜᐟဏញܫનȐcorticotrophin-releasing hormone, CRHȑޑઓϡՏܭ ΠຎЫ࠻ਡȐparaventricular nucleusȑޑǹѦࣚޑڈᐟࢂڀ࠶ુ܄ޑǴ
ૻဦҗགᏔ۔ډΠຎЫǴڈᐟځϩݜᐟဏញܫનǹ၀ញܫનၸ တࠟဏߐેسډၲတࠟဏယǴ٬ځϩݜᐟဏҜ፦નȐadrenocorticotropic hormone, ACTHȑǴ೭٤ဏҜ፦નӆѐڈᐟဏҜ፦ӝԋဏ ᗐҜ፦ᐟનȐglucocorticoidȑ (Engelmann, Landgraf, & Wotjak, 2004)Ƕ೭٤ᐟન
ញܫډՈనύၲډӄيǴჹނӧय़ჹᆙ࡚٣ҹਔࢂख़ाޑǶނࡋၸᆙ
࡚٣ҹࡕǴॄӣ㎸ፓᐒڋ௴٠ڋΠຎЫ-တࠟᡏ-Ҝ፦ືጕޑࢲǴٰ
ᗉխՈనύޑဏᗐҜ፦ᐟન֖ໆၸଯ (Sapolsky, Romero, & Munck, 2000)Ƕ ӢԜǴໆෳҜ፦✉Ȑcorticosteroneȑ܈Ҝ፦ᎇȐcortisolȑ೭٤ဏᗐҜ፦ᐟન ޑᡂϯෳໆނޑᓸΚೀϐख़ाғࡰ (M÷stl & Palme, 2002)ǶԖࣴ
زࡰрǴ૨ηװᔐޑҢጄޣǴځՈనύޑҜ፦✉֖ໆϲǹԶӧᢀჸҢጄ ޣᎁၶװᔐޑᢀჸޣǴՈనύҜ፦✉֖ໆΨϲ (Kavaliers et al., 2003)ǴᡉҢ Αय़ჹڀ࠶ુޑڈᐟނόѝࢂԋঁᡏౢғᓸΚϸᔈǴᢀჸӕᜪᎁၶڀ࠶ુ܄ޑ ٣ҹΨ٬ᢀჸޣౢғᓸΚϸᔈǶԖࣴزҭࡰрǴ܄ঊႵӧᆶځѬӕᜪՉ
ԛᝡݾ܄ϕΠ௳ǴՈనϣޑҜ፦ᎇ֖ໆϲ (Huhman, Moore, Ferris, Mougey, & Meyerhoff, 1991)ǶՠӧڀԖ௳ᡍΠѐᢀჸӕᜪՉӕኬ܄፦ޑ ϕ٣ҹǴࢂցԋᢀჸޣޑҜ፦ᎇ֖ໆӢԜڙډቹៜǴ߾ࢂঁ҂ޕኧǴᗨฅ ԖࣴزࡰрವΟКլαჯങ ( Oreochromis mossambicus ) ӧؒԖҺՖᡍΠѐ ӧᢀჸӕՔޑᝡݾ܄ϕ٣ҹਔǴᡏϣޑ 11-keto-࿐ڰ✉Ȑ11-keto-testosteroneȑ
ک࿐ڰ✉ޑ֖ໆϲଯǴᡉҢᢀჸޣჹᝡݾ܄ϕ٣ҹՉᢀჸǴғࢂڙ ډቹៜޑ (Oliveira, Lopes, Carneiro, & Canario, 2001)Ƕ
ဏᗐҜ፦ᐟનޑኬҁڗளٰྍЬाёаϩԋߟΕ܄ᆶߚߟΕ܄ޑБ ԄǶӧߟΕ܄БԄǴЬाࢂၸ׀ᓉેǵᆂǵЈ᠌ऀڈ܈ޣᘐᓐٰ௦ளՈన ฅࡕՉϩǹӧߚߟΕ܄БԄǴ߾ࢂԏނޑᕨߡٰෳໆᕨߡϣϐဏ ᗐҜ፦ᐟનжᖴނ֖ໆǶ࣬ၨϐΠǴߚߟΕ܄БԄКၨ໔ௗЪኬҁᇆΚ
ਔǴࢂҞКၨόදၹՠΞКၨҔܭՉᓸΚ࣬ᜢࣴزޑБԄǴচӢࢂߟΕ
܄௦ՈБԄԖځᝄख़ޑલᗺᆶज़ڋǴٯӵǺӧՉ௦Ոਔሡज़ڋނޑՉ܈ޣ
ՉഞᎈǴՠӵԜբёૈ٬ளՈనύဏᗐҜ፦ᐟન֖ໆቚуԶᏤठჴᡍ ਏ݀ԡࢉ (Pace & Spencer, 2005; Vahl et al., 2005)Ǵӵ݀ࣴزޣௗΠٰᗋགྷाᢀ ჸނӧځѬՉࣁբޑ߄ǴՉ௦Ոਔ܌ԋޑᓸΚёૈ൩ቹៜނޑ Չࣁ߄ǹќѦǴࣴزޣӵ݀གྷाᢀჸڙ၂ޣՈనύဏᗐҜ፦ᐟન֖ໆޑೱ
ុਔ໔ᗺᡂϯǴനதـޑբݤࢂ٬Ҕόӕಔޑڙ၂ޣӧόӕਔ໔ᗺѐՉ௦ՈǴ ՠ೭ኬޑբݤሡा٬Ҕεໆڙ၂ޣЪคݤஒᅟᆾᡂϯޑঁᡏৡ౦܄๏Ե ቾѐǶќБय़Ǵ٬ҔߚߟΕ܄БԄᗨёှ،ॊޑલᗺϷज़ڋǴՠሡԵቾӵ Ֆᒧᕨߡԏޑਔ໔ᗺǴӢࣁჴሞჴᡍᏹϐਔ໔ᗺᆶᅟᆾжᖴނӧᕨߡ ϣୀෳډϐਔ໔ᗺϐ໔Ԗࢤۯ᪓ਔ໔Ǵ܌аӵՖᒧᕨߡԏޑਔ໔ᗺǴ
،ۓࣴزޣࢂցૈୀෳډჴᡍᏹ܌ԋޑᓸΚϸᔈϐਏ݀ǶࣴزวǴӧ
ނيݙ 3H-Ҝ፦✉Ȑ3H-corticosteroneȑǴݙਔ໔ࢂวғӧλႵȐC57BL/6Jȑ ޑསຼයаϷߝຼයϐਔǴ܌ෳໆډޑۯ᪓ਔ໔ϩձࢂ 4 λਔаϷ 10 λਔѰѓ (Touma, Sachser, Möstl, & Palme, 2003)ǹՠӧεႵȐSprague-Dawley ratȑي߾
ࣁ 17 λਔѰѓ (Bamberg, Palme, & Meingassner, 2001)ǴᡉҢΑۯ᪓ਔ໔ޑᡂ౦
܄ࢂࡐεޑǴڙჴᡍࢂวғӧϺύޑবঁຼයύаϷ܌٬ҔޑނࠔسԶԖ
܌ቹៜǶନԜϐѦǴ၀ࠔᅿޑᓸΚϸᔈૈցၸ೭ᅿߚߟΕ܄מೌٰୀෳрǴΨ
ࢂ٬ҔԜБԄਔሡाԵቾޑ (Rupert, 2005)Ƕᆕӝа܌ॊǴךॺᇡࣁǴҗܭࣴ
زޗ܄ૻ৲᠍ڗޑᚒǴሡाᡣ܄ঊႵௗڙೱុޑᢀჸᏢಞ૽ግаϷᏢಞਏ
݀ෳ၂ǴӢԜ೭ኬޑᐕำёаႣයុࢤόอޑਔ໔ǴӢԜӧόቹៜނܭ ೭ࢤਔ໔ϣޑᏢಞᐕำаϷނޑВதࢲΠǴ௦ҔߚߟΕ܄Бݤѐޗ܄
ૻ৲᠍ڗᐕำޑՉύ܌ౢғޑᓸΚϸᔈёૈࢂၨӝޑǶ
ມǵਡЈୢᚒᆶှ،ౣ
ҁፕЎύԖΟঁЬाޑਡЈୢᚒǴਥᏵ೭٤ୢᚒǴךॺीΑѤঁჴᡍǺ
ǵࡌҥคঁᡏޗҬᡍచҹΠޑޗ܄ૻ৲᠍ڗϐ௶ճ٥܄ঊႵኳࠠ
ၸჹޗ܄ૻ৲᠍ڗᚒޑӣ៝ǴךॺΑှډǴᢀჸޣჹӕᜪ܌วғޑ ᝡݾ܄ϕ٣ҹՉȸᢀჸȹǴԶᢀჸޣவ٣ҹύಞளȸૻ৲ȹǴ٠ஒૻ৲٬
ҔӧࡕុᏯରӝϐύǴࣁϩڀόӕᏯରૈΚޑჹЋϐ٩ᏵǶӢԜǴӧךॺ
܌ࡌҥޑ௶ճ٥܄ঊႵኳࠠύǴךॺሡाܴᢀჸޣԖӧჹڬᎁޑӕᜪϕ٣ ҹՉᢀჸǴ٠வύಞளૻ৲Ƕ܌аӧჴᡍǴךॺᡣؒԖҺՖѺରᡍޑ܄
ڙ၂ޣѐᢀჸǺќѦٿଫ܄ҢጄޣՉѺରǹ܈ޣ႖៓ᆛՉύ܄ޗҬϕ
Ǵ೭ٿᅿόӕ܄፦ޑ٣ҹǶฅࡕܭќঁୱύज़ڋҢጄޣޑՉǴаԛග ٮঁۓҢጄޣښၰޑБԄǴෳໆڙ၂ޣჹۓښၰޑϸᔈǶӵԜՉࣁෳໆБ ԄǴ٬ளךॺૈෳໆډڙ၂ޣ٩ၸѐჹۓښၰޑᏫ܌߄рޑՉࣁϸᔈǶ ӧ೭ኬޑीϐΠǴךॺଷڙ၂ޣӵ݀ԖჹҢጄޣ܌ޑՉՉᢀჸǴ߾ό ӕ܄፦ޑϕ٣ҹ܈ޣόӕ܄፦٣ҹύޑҢጄޣǴ٬ளڙ၂ޣӧᢀჸΠޑ܈
ޣࡕុ༘ᆪډۓҢጄޣښၰϐਔޑՉࣁ߄Ԗ܌όӕǶќѦǴଷڙ၂ޣԖ வᝡݾ܄ϕ٣ҹύಞளૻ৲Ǵ߾ڙ၂ޣ༘ᆪډϕ٣ҹύޑគৎҢጄޣϐښၰ
܌߄рޑՉࣁόӕܭ༘ᆪډϕ٣ҹύޑᒡৎҢጄޣϐښၰ܌߄рޑՉ ࣁǶ
ΒǵঁᡏޗҬ௳ᡍࡕޑޗ܄ૻ৲᠍ڗ
ၸჹᢀჸᏢಞᚒᚒޑӣ៝ǴךॺΑှډǴঁᡏӃޑॄӛᡍёૈቹ
ៜᢀჸޑᏢಞᐕำаϷࡕុޑᏢಞ߄ǴӢԜӧჴᡍΒǴךॺӧჴᡍ܌ࡌҥ рޑޗ܄ૻ৲᠍ڗϐჴᡍᏹբՉࣁኳࠠύуΕঁᡏޑޗҬ௳ᡍǴᡣڙ၂ޣ ӧڀԖԛ௳ᡍΠѐௗڙӕኬޑᢀჸᏢಞ૽ግаϷࡕុޑՉࣁෳ၂Б ԄǶ௳ڙ၂ޣޑჹЋᆶڙ၂ޣஒाᢀჸډޑҢጄޣࢂόӕޑڈᐟނǴ೭ኬޑ
ीБԄஒ٬ளڙ၂ޣӧࡕុޑᏢಞ߄ǴځϸᔈଞჹޑஒࢂӃᢀჸډޑ ϕ٣ҹύޑۓҢጄޣǴԶߚଞჹӃ๏ϒ௳ᡍޑჹЋǶ
Οǵޗ܄ૻ৲᠍ڗᐕำΠᆶࡕុޑૻ৲٬ҔਔޑᓸΚϸᔈ
ӧჴᡍΟύǴЬाࢂໆෳ܄ঊႵӧҁፕЎޑჴᡍᏹΠϐᓸΚϸᔈǴёϩ ԋჴᡍΟ A ᆶჴᡍΟ BǶࣁΑໆෳ೭ኬޑᓸΚϸᔈǴךॺςߚߟΕ܄БԄǴٰ
ୀෳڙ၂ޣޑᕨߡϣҜ፦ᎇжᖴނȐfecal cortisol metabolites, FCMȑ֖ໆǶךॺ ςΑှډӧ٬ҔߚߟΕ܄БݤਔǴሡԵቾ܌ᒧޑᕨߡԏਔ໔ᗺǴӢࣁѬஒ،
ۓࢂցૈໆෳډނӢჴᡍᏹ܌ౢғޑᓸΚϸᔈǴҗܭၸѐӕኬаߚߟΕБݤ ӧ௶ճ٥ঊႵيᓸΚ࣬ᜢᚒޑࣴزѝԖጇǴ၀ࣴز௦ڗԏ٣ҹࡕ Ӛ 24 λਔϣڙ၂ޣ܌௨ܫрٰޑᕨߡ (Chelini, Souza, Cortopassi, Felippe, &
Oliveira, 2006)ǴځѬޑࣴز߾ӭаλႵ܈εႵࣁڙ၂ޣ (Bamberg et al., 2001;
Touma et al., 2003)ǶҗܭঊႵޑᡏࠠϟΕλႵᆶεႵϐ໔Ǵՠ߈՟ܭλႵǴࣁΑ ዴߥ܌ᒧਔ໔ᗺૈୀෳډᓸΚϸᔈǴךॺ٩ჴᡍҞޑሡǴ௦ڗ 3 ঁλਔ ൩ԏԛ܄ঊႵ܌௨ܫрٰޑᕨߡ܈ޣ 18 λਔϣ܌௨ܫрٰޑᕨߡǶࣁ ΑዴᇡҁፕЎύ܌٬ҔޑߚߟΕ܄ᕨߡϣҜ፦ᎇжᖴނୀෳݤૈໆෳډڙ၂ޣ ޑᓸΚϸᔈǴךॺӧჴᡍΟ AǴ٬ҔΑԜמೌѐໆෳ܄ঊႵӧڀԖόӕ܄፦ޑ
ঁᡏޗҬᡍΠޑᓸΚϸᔈǴӧ೭ኬޑޗҬᡍΠǴڙ၂ޣڀԖԛ௳ޑ
ᡍ܈ޣԛᏯയќଫ܄ঊႵޑᡍǶךॺෳǴӵ݀೭ኬޑמೌૈᔈҔӧ
܄ঊႵي٠ໆෳрځ܌ౢғޑᓸΚϸᔈǴ߾ၸѐၸߟΕ܄௦ՈБݤӧ܄
ঊႵيаᜪ՟ޑᏹ܌ளډޑ่݀ (Huhman et al., 1991)ǴΨૈӧჴᡍΟ A ύ аߚߟΕמೌٰளډᡍǶӧჴᡍΒޑीΠǴךॺᡣڙ၂ޣᆶќଫ܄ঊႵ
Չᝡݾ܄ϕԶᏱԖ௳ޑᡍǴӆᡣڙ၂ޣаᢀޣޑيϩѐय़ჹឦܭӕ
ኬ܄፦ޑᝡݾ܄ϕ٣ҹޑวғǴךॺෳǴڙ၂ޣӧ೭ኬޑᏢಞᐕำύаϷࡕ
ុ༘ᆪډϕ٣ҹύޑគৎҢጄޣϐښၰਔǴࡐёૈࢂೀܭԖᓸΚޑރᄊǶӢ ԜӧჴᡍΟ BǴךॺճҔԜמೌѐୀෳჴᡍΒύޑڙ၂ޣϐᓸΚϸᔈǶ
ಃΒകʳ ჴᡍᆶБݤ
ಃʳ ჴᡍ
ҁჴᡍޑҞޑࢂճҔঊႵڀԖϐሦୱװᔐՉࣁࣁኳԄǴᡣؒԖҺՖޗҬᡍ ޑԋԃ܄ঊႵѐᢀჸќѦٿଫ܄ঊႵޑޗҬϕՉࣁǴᙖԜٰࣴزޗ܄ૻ
৲᠍ڗޑຝࢂցӸӧܭঊႵيǴΨ൩ࢂ҂ԖޗҬᡍޑᢀჸޣૈցӧᢀჸٿ ଫҢጄޣޑѺର٣ҹࡕǴჹ೭ٿଫҢጄޣ߄рόӕޑՉࣁǴᙖаፕޗ܄ૻ
৲᠍ڗӵԜޑᢀჸᏢಞБԄǴӧԋԃঊႵيޑዴӸӧǶନԜϐѦǴၸғࡰ
ٰᡍᢀჸၸำύ܌ԋޑቹៜǶ
ǵڙ၂ޣ
ҁჴᡍ௦Ҕ܄௶ճ٥ঊႵȐMesocricetus auratusȑǴрғ 8 ຼࡕԾ୯ৎჴ ᡍނύЈᖼளǴаঁձႩᎦ᠈Ȑᆫ✊༟ጤ፦Ǵߏ 33ǵቨ 21ǵଯ 15 Ϧϩȑࣁ ႖ᚆൂՏޑ IVC سȐindividual ventilated caging systemȑႩᎦܭ୯ҥѠεᏢ Јسނ܊ϣǴ᠈ϣᎎҏԯఒპǴ٠кϩٮᔈႩᆶНǶჴᡍύ܌٬Ҕ ڙ၂ޣӅ 34 ଫǴႵសऊࣁ 16 ډ 20 ຼǶނ܊ϣۓࣁ 12Ǻ12 ॹᙯϐВڹຼයǴ སຼයவԐ 8 ᗺ໒ۈǴྕࡋᆢ 22 ± 2ʚǴྒྷࡋᆢӧ 50 %ѰѓǶ
ΒǵՉࣁໆෳᏔ
ჴᡍύ܌٬ҔޑՉࣁໆෳᏔӅԖ 2 ᅿǴϩձࣁᢀჸᏢಞጃȐobservational learning boxȑᆶ U ࢭȐU mazeȑǶ
() ᢀჸᏢಞጃ
٬ҔᢀჸᏢಞጃޑҞޑࢂբࣁᢀჸᏢಞ໘ࢤ૽ግϐҔǴᡣڙ၂ޣёаᢀჸќ Ѧٿଫ܄ঊႵޑϕǶӵკ A ܌ҢǴᢀჸᏢಞጃȐߏ 44 Ϧϩǵቨ 44 Ϧϩǵ ଯ 15 ϦϩȑҗᓸլΚ፦܌ᇙԋǴБԖܴᓸլΚ݈ᙟᇂǴБߡჴᡍޣޑᢀ ჸǴ٠ᗉխނޑಥǹځϣճҔᓸլΚ݈ᆶқՅλ៓᠈Ȑߏ 8.5 Ϧϩǵቨ 8.5
Ϧϩǵଯ 13.5 Ϧϩȑஒނ۶Ԝ႖໒ٰǶᢀჸᏢಞጃёаϩࣁȨᢀჸȩǵȨϕ
ȩᆶȨࡑȩΟঁϩǶᢀჸࢂҗٿঁқՅλ៓᠈η٠௨܌ಔԋǴٿঁ
᠈ηύ໔ақՅόܴ႖݈႖໒ٰǴ٬ளٿଫڙ၂ޣ໔۶Ԝคݤ࣮ډჹБǴ
ԛჴᡍԿӭٿଫڙ၂ޣёаϩձܫΕλ៓᠈ύӕਔՉᢀჸᏢಞǶϕࣁ
ߏБࠠȐߏ 22 Ϧϩǵቨ 15 Ϧϩǵଯ 15 ϦϩȑǴёᡣٿଫ܄ҢጄޣӧԜϣ
ՉϕǶࡑࣁΟفࠠޜ໔Ȑᜐ 15 Ϧϩǵۭᜐ 8.5 Ϧϩǵ௹ᜐ 17 Ϧϩǵ ଯ 15 ϦϩȑǴࢂᢀჸᏢಞ૽ግᗋ҂໒ۈǴҢጄޣޑኩࡑϐೀǶᢀჸᏢಞጃϣӚ
ޑ࣬ჹՏǺᆶᢀჸ࣬ᎃޑࢂϕǴٿঁ໔аλ៓᠈ηҁيޑ៓ํᆛஒ ٿ႖໒ǹӧϕٿޑࢂࡑǴނࢂҗٿޑࡑΕϕǴࡑ
ᆶϕ໔߾аё౽ԄқՅόܴᓸլΚ݈႖໒ǶϕޑѦൎᕅᏛᆶ
ᢀჸޑѦൎᕅᏛӕኬڀϾࢰȐϾ৩ελࣁ 0.7 ϦϩȑǴаճࢬޑࢬ೯Ƕӧ ᢀჸࡕБޑᕅѦǴ߾းԖႝܜ॥৻ǴܭჴᡍՉύ໒௴Ǵࢬஒҗϕޑ ѦൎᕅᏛΕϕനࡕډၲᢀჸǹ೭ኬޑीஒ٬ளښၰޑሀࢂൂӛޑǴ ନΑ٬ڙ၂ޣܰᆪډҢጄޣޑښၰѦǴᗋૈᡣҢጄޣόܰᆪډڙ၂ޣޑښၰԶ
ૈݙܭӧ҅ՉޑޗҬϕǴӢԜҢጄޣஒόܰӢڙ၂ޣޑӸӧԶቹៜ ΑځՉࣁ߄Ƕ
--- კ A ܭԜ --- (Β) Uࢭ
٬Ҕ U ࢭޑЬाҞޑࢂҔٰෳ၂ڙ၂ޣჹܭ҂ٰஒᢀჸډ܈ࢂςᢀ ჸၸϕޑҢጄޣӧќঁόӕӦύϐϸᔈǶࢭЬा፦ࣁᓸլΚǴۭࣁ қՅǴନΑࢭύҔٰϩ႖όӕୱޑ౽Ԅ႖݈ࣁқՅѦǴࢭޑᕅᆶϺ݈
ࢂܴޑǴаճჴᡍޣᢀჸނޑࢲރݩǶӵკΒ A ܌ҢǴU ࢭޑѰ ѓٿᆄϩձҗঁ V ࠠᖉ܌ಔԋǴঁ V ࠠᖉޑٿᖉ໔֨فࣁ 120±ȐٿᖉӚࣁ ߏ 42 Ϧϩǵቨ 10 Ϧϩǵଯ 15 ϦϩȑǴঁ V ࠠᖉഗᆄӚԖঁڈᐟނǴҗ
қՅλ៓᠈ηȐߏ 8.5 Ϧϩǵቨ 8.5 Ϧϩǵଯ 13.5 Ϧϩȑ܌ԋǶӧ V ࠠٿᖉۭ
ᆄӚΕТё౽Ԅක݈ȐόܴԖऀϾޑᐋ✊࣒ዟ႖݈ǹࠆ 0.3 ϦϩǴϾ৩ ελࣁ 0.7 ϦϩȑǴԜٿТක݈ᆶࢭۭᜐޑڰۓԄᕅᏛȐόܴԖऀϾޑᐋ✊
࣒ዟ႖݈ǴϾ৩ελࣁ 0.7 Ϧϩȑԋ҅Οفޑࡑ ȐᜐߏӚࣁ 10 Ϧ ϩȑǶӧ U ࢭύǴࡑϐё౽Ԅක݈౽໒ਔǴନΑڈᐟނѦǴڙ ၂ޣёаᒧΕࢭޑҺୱǶࣁΑБߡᒵڙ၂ޣӧ U ࢭύޑՏǴ
ࢭޑ V ࠠѰѓᖉӚёϩԋϖঁୱǴԶ V ࠠѰѓᖉё٩ᏵڈᐟނޑԖคϩձ
ڮӜࣁȨڀڈᐟނޑٗᖉȩᆶȨόڀڈᐟނޑٗᖉȩǴঁ V ࠠᖉΞёӆ ಒϩԋȨ߈ࡑޑࡑȩᆶȨ߈ڈᐟނޑڈᐟނȩǴ೭ٿঁୱΞё аਥᏵຯᚆᇻ߈ӆ֡ϩԋȨ߈ࡑȩȐߏ 21 ϦϩȑᆶȨᇻࡑȩȐߏ 21ϦϩȑаϷȨᇻڈᐟނȩȐߏ 21 ϦϩȑᆶȨ߈ڈᐟނȩȐߏ 21 Ϧϩȑǹନ ԜϐѦǴӧ߈ڈᐟނޑഗᆄຯᚆёаௗ܈༘ᆪډڈᐟނ 2 ϦϩϣޑୱǴ
ᆀࣁȨ߈ڈᐟނޑനᆄȩǶӢԜ U ࢭύঁ V ࠠᖉϣޑ೭ϖঁୱவ ڈᐟނଆǴϩձࣁ߈ڈᐟނޑനᆄǵ߈ڈᐟނǵᇻڈᐟނǵᇻࡑǵ аϷ߈ࡑǶࣁΑᙁϯჴᡍኧᏵБߡှଆـǴךॺஒȨ߈ڈᐟނޑനᆄȩ уȨ߈ڈᐟނȩаϷȨᇻڈᐟނȩ೭ΟঁୱӝᆀࣁȨښၰගٮȩǶ
ঁ U ࢭޑीࢂ௦ڗஏഈՠё೯॥ޑБԄǴၸܫܭࢭۭᜐᕅᏛࡕБ ޑႝܜ॥৻ǴܭჴᡍՉύ໒௴ǴࢭѦޑޜஒҗڈᐟނܜΕٿᖉനࡕ ډၲࢭۭᜐࡕ௨рǶ
--- კΒ A ܭԜ --- ΟǵՉࣁբᆶࡼෳำׇ
ҁჴᡍՉޑՉࣁբࣁȨᢀჸᏢಞޑᏹȩǴҞޑࢂᡣ܄ঊႵѐᢀჸќ Ѧٿଫ܄ঊႵޑޗҬϕՉࣁǴᙖаࣴزޗ܄ૻ৲᠍ڗǶӧ໒ۈՉՉࣁჴ ᡍǴջҗჴᡍޣϺڗঊႵ٠ગख़ԿϿຼǴ٬ځಞᄍჴᡍޣޑᏹբǶՉࣁ
բ֡ӧނ܊ϐঊႵՉࣁෳ၂࠻ϣՉǴନΑϺ݈ޑЍआՅВӀᐩѦǴ ᗋуޑϤΜґᘚᐩ༾১ӀጕࣁྣܴȐྣܴࡋࣁ 30 LuxȑǶ܌ԖՉࣁᏹ
ᆶෳ၂֡Ծύϱ 12 ᗺၸࡕ໒ۈՉǴЪӧఁΎᗺа่״ǶӧԜբύǴ ڙ၂ޣܭᢀჸጃаϷ U ࢭύޑ܌ԖՉࣁ߄ࣣаឪቹᐒȐJVCȑᒵΠٰǴ Զނޑ܌ԖՉࣁ߄֡аዸᒮ܈ឲᒵӧࠠႝတύޑՉࣁᒵ೬ᡏ Ethom
ٰᒵ (Shih & Mok, 2000)ǶӵკΟ܌ҢǴՉࣁჴᡍᆶᏹբำׇӅೱុՉ 6 ϺǴӧ೭ 6 Ϻύޑᏹёа٩ׇϩԋѤঁ໘ࢤǴϩձࣁಃϺޑȨಞᄍϯ໘ࢤȩ Ȑhabituation phaseȑǵಃΒϺޑȨෳ໘ࢤȩȐpretest phaseȑǵಃΟډಃϖϺъ ࢤޑȨᢀჸᏢಞ໘ࢤȩȐobservational learning phaseȑǵᆶಃϖϺࡕъࢤډಃϤϺ ޑȨᏫෳ၂໘ࢤȩȐmemory testing phaseȑǶ೭Ѥঁ໘ࢤޑ၁ಒᏹำׇϩॊӵ ΠǺ
--- კΟܭԜ --- () ಞᄍϯ໘ࢤ
Ԝ໘ࢤޑҞޑӧܭफ़եڙ၂ޣჹढ़ғᕉნޑขቾǴ٠٬ځዕՉࣁໆෳᏔ
ǶӧϤϺჴᡍࢬำޑಃϺՉಞᄍϯ໘ࢤǴ܌Ԗޑڙ၂ޣࣣϩձΕς
٬Ҕ 70 %ޑଚᆒᔔଳృޑ U ࢭύǴҺځԾҗ 10 ϩដǶନΑ U
ࢭѦǴڙ၂ޣࡕុΨΕқՅλ៓᠈ηύǴܫΕᢀჸᏢಞጃϣޑᢀჸՉ 30ϩដޑಞᄍϯǶ่״ࡕǴஒڙ၂ޣӼӣচۚՐႵ᠈ϣǶ
(Β) ෳ໘ࢤ
Ԝ໘ࢤޑҞޑࢂዴᇡஒΕᢀჸᏢಞ໘ࢤޑڙ၂ޣჹܭ҂ٰाᢀჸޑҢጄ ޣڀԖۓำࡋޑᑫ፪ǶӧϤϺჴᡍࢬำޑಃΒϺՉ䬲໘ࢤǴ܌Ԗޑڙ၂ ޣࣣϩձΕς٬Ҕ 70 %ޑଚᆒᔔଳృޑ U ࢭύٰՉෳ၂ǴԜ໘ࢤ х֖ٿঁ 3 ϩដޑೱុ၂ǴϩձࣁȨڙ၂ޣୃӳՏෳ၂၂ȩᆶȨय़ჹڈᐟ ނޑᄊࡋෳ၂၂ȩǶӧڙ၂ޣୃӳՏෳ၂၂ύǴ٠คڈᐟނޑӸӧǹय़ჹ
ڈᐟނޑᄊࡋෳ၂၂ύ܌٬ҔޑڈᐟނǴ߾ࢂڙ၂ޣ҂ٰӧᢀჸᏢಞ໘ࢤύ܌
࣮ډޑٿଫҢጄޣύޑځύଫǶڙ၂ޣୃӳՏෳ၂၂ޑҞޑӧܭᒧрௗΠ
ٰाՉޑय़ჹڈᐟނޑᄊࡋෳ၂၂ύǴڈᐟނाܫܭ U ࢭޑѰᜐ܈
ѓᜐޑՏǶ໒ۈՉԜ၂ਔǴڙ၂ޣܫܭࡑϣǴӧ॥৻໒௴ 30
ࣾࡕǴஒё౽Ԅක݈ܜ໒Ǵᡣڙ၂ޣӧ U ࢭϣԾҗǶҗܭ U ࢭ Ѱѓ V ࠠᖉӚჄϩр 5 ঁୱǴӢԜ၂ύڙ၂ޣӧ೭Ӆ 10 ޑӚୱϣϐ
੮ਔ໔аϷӚୱϐऀຫԛኧჴᡍޣᒵΠٰǴڙ၂ޣӧӚୱޑऀຫԛ ኧຎࣁڙ၂ޣӧӚୱϐࢲΚޑࡰǶӧԜ၂܌ໆෳޑڙ၂ޣϐՉࣁࡰ
ࣁȨU ࢭϐࢲΚȩǴջڙ၂ޣӧ U ࢭϣޑ 10 ঁୱ໔ޑᕴऀຫԛኧǶ
3 ϩដޑਔ໔ډၲਔǴஒ॥৻ᜢǴڙ၂ޣ߾ܫӣࡑύǴࡑΠ
ঁय़ჹڈᐟނޑᄊࡋෳ၂၂Ƕ
ڙ၂ޣӧय़ჹڈᐟނޑᄊࡋෳ၂၂܌ाय़ჹޑڈᐟނࢂ௦ڗᒿᐒϩ ࢴǴӧڈᐟނޑᒧǴࣁΑڋڙ၂ޣ҂ٰ܌ᢀჸډҢጄޣ۶Ԝ໔ޑѺରࠔ፦
ᆶയॄǴךॺ٣Ӄ૽ግ٠ࡷᒧрץڀԖೱՍᕇയᡍޑគৎЪၨڀமװᔐ
܄ޑץװᔐޣȐfighterȑǴߐҔٰѺॹჹЋǹӕਔΨྗഢΑץڀԖऩυѨ
௳ᡍЪၨόڀװᔐૈΚޑᒡৎǴ٬ځӧय़ჹڀװᔐ܄ޑჹЋ೯தѝԖѺޑ ҽǶӧෳ໘ࢤ໒ۈՉǴךॺᙣޑᡣӚঁҢጄޣᆶځѬӕᜪՉၸӭ ԛޑޗҬϕǴٰዴߥځ܌ڀيҽᗋࢂᆢόᡂޑǶय़ჹڈᐟނޑᄊࡋෳ၂၂ ޑᏹำׇӵӕڙ၂ޣୃӳՏෳ၂၂ኬǴڈᐟނ܌ᘍܫޑՏࢂҗڙ၂ޣ ӧՉڙ၂ޣୃӳՏෳ၂၂ਔǴࡑӧ U ࢭٿᖉύځύᖉϣຯᚆڈᐟ ނന߈ϐȨ߈ڈᐟނޑനᆄȩуȨ߈ڈᐟނȩ೭ٿޑᕴਔ໔ၨߏ ޣࣁڈᐟނܫޑᆄǶӕኬᒵ၂ύڙ၂ޣӧӚୱϐऀຫԛኧᆶ੮ਔ ໔ǶӧԜ၂ाໆෳޑڙ၂ޣϐՉࣁࡰӅԖѤঁǴϩձࣁڀڈᐟނޑٗᖉ ϐȨډၲ߈ڈᐟނޑനᆄϐ܌ሡਔ໔ȩᆶȨښၰගٮϐ੮ਔ໔ȩǹȨόڀ ڈᐟނޑٗᖉϐࢲΚȩǵᆶȨόڀڈᐟނޑٗᖉϐ੮ਔ໔ȩǶ၂่״
ࡕǴஒڙ၂ޣӼӣচۚՐႵ᠈ϣǴU ࢭ߾٬Ҕ 70 %ޑଚᆒᔔଳృᗉխ
੮ΠҺՖښǶ
ࣁΑዴᇡڙ၂ޣჹڈᐟނڀԖۓำࡋޑᑫ፪ᆶݙཀำࡋǴךॺۓΠΑᑔᒧ
ྗࣁǺӧय़ჹڈᐟނޑᄊࡋෳ၂၂ޑ 3 ϩដϣǴѸ੮ӧڀڈᐟނޑٗ
ᖉຯᚆڈᐟނന߈ϐȨ߈ڈᐟނޑനᆄȩуȨ߈ڈᐟނȩ೭ٿঁ
ୱޑਔ໔ᕴکεܭ 60 ࣾаǴ܈ޣࡑӧȨ߈ڈᐟނޑനᆄȩޑਔ໔Ԗ 40
ࣾаǶ೯ၸෳ໘ࢤᑔᒧྗޑڙ၂ޣஒᒿᐒϩࢴډΟಔǴϩձࣁȨय़ჹύ
܄ڈᐟಔȩǵȨय़ჹᒡৎڈᐟಔȩǵᆶȨय़ჹគৎڈᐟಔȩǶӧֹԋ U ࢭޑ
ෳࡕǴ೭٤ڙ၂ޣӆԛܫΕᢀჸᏢಞጃϣޑᢀჸՉ 30 ϩដޑಞᄍ ϯǴ٬ځкϩ٠ಞᄍᢀჸаϷλ៓᠈Ƕ
(Ο) ᢀჸᏢಞ໘ࢤ
Ԝ໘ࢤޑҞޑࢂᡣڙ၂ޣೱុΟϺᢀჸځѬٿଫ܄ঊႵวғޗҬϕ٣ ҹǴ٠ၸᢀჸ೭ኬޑϕ٣ҹٰᏢಞϩ܌ᢀჸډޑҢጄޣϐ໔ޑόӕǶӧϤ ϺჴᡍࢬำޑಃΟϺ໒ۈՉೱុΟϺޑᢀჸᏢಞ໘ࢤǴڙ၂ޣᒿᐒϩࣁჴ ᡍಔᆶڋಔǴჴᡍಔᢀჸډٿଫ܄ঊႵޔௗϕ٠วғѺର٣ҹǴᆀϐࣁ ȨᢀჸѺର٣ҹಔȩǹڋಔ߾ᢀჸٿଫ܄ঊႵ႖λ៓᠈໔ௗϕՠคݤว ғѺର٣ҹǴᆀϐࣁȨᢀჸύ܄٣ҹಔȩǶӵკѤ܌ҢǴӧᢀჸύ܄٣ҹಔύǴ ךॺஒځύଫҢጄޣᜢΕλ៓᠈ηύǴܭϕޑύѧǴԶᡣќѦଫҢጄ ޣࡑӧϕύޑ៓᠈ηѦय़Ǵ٬ځᆶ៓᠈ηϣޑҢጄޣՉ໔ௗ܄ϕ࣬ޑ ՉࣁՠόวғѺରՉࣁǶӃ೯ၸෳ໘ࢤᑔᒧޑڙ၂ޣǴਥᏵځӧ၀໘ࢤਔ
܌य़ჹޑڈᐟނϐيҽङඳǴԶஒڙ၂ޣᒿᐒϩࢴډᢀჸѺର٣ҹಔ܈ᢀჸύ
܄٣ҹಔǶКӵӧෳ໘ࢤਔǴӧჴᡍޣޑᒿᐒϩࢴΠǴӵ݀ڙ၂ޣय़ჹޑڈᐟ ނޑيҽङඳࣁᒡৎޑҢጄޣǴ߾ᢀჸᏢಞ໘ࢤਔǴڙ၂ޣ൩ϩࢴډᢀჸѺ ର٣ҹಔύǴӵԜڙ၂ޣωૈᢀჸډيҽङඳࢂᒡৎޑҢጄޣќଫគৎҢጄ ޣ๏Ѻ௳ǹӵ݀ڙ၂ޣय़ჹޑڈᐟނيҽङඳࢂគৎǴ߾ᢀჸᏢಞ໘ࢤਔǴڙ၂ ޣஒᒿᐒϩࢴډᢀჸύ܄٣ҹಔ܈ޣᢀჸѺର٣ҹಔϐύǴӵԜڙ၂ޣஒᢀ ჸډԜڈᐟނيҽङඳࣁគৎޑҢጄޣǴܭύ܄٣ҹύᆶќଫҢጄޣՉύ
܄ϕǴ܈ܭѺର٣ҹύѺ௳ќଫᒡৎҢጄޣǶӢԜǴय़ჹύ܄ڈᐟಔȐn = 13ȑ ޑڙ၂ޣᢀჸޑࢂٿଫҢጄޣՉύ܄ϕǴջࣁᢀჸύ܄٣ҹಔȐn = 13ȑǹԶ य़ჹᒡৎڈᐟಔȐn = 10ȑᆶय़ჹគৎڈᐟಔȐn = 11ȑύޑڙ၂ޣӧΟϺޑᢀჸ Ꮲಞ໘ࢤӕኬᢀჸٿଫҢጄޣՉѺରϕǴջࣁᢀჸѺର٣ҹಔȐn = 21ȑǶ
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ಃΟϺՉᢀჸᏢಞ໘ࢤޑಃϺ૽ግǴӧ૽ግ໒ۈஒόܴό
ޑᓸլΚ݈ߔ႖ӧᢀჸᆶϕϐ໔Ǵ٬ڙ၂ޣӧΕᢀჸࡕኩਔ࣮όډϕ
ϣޑҺՖࢲǶڙ၂ޣܭᢀჸࡕǴٿଫջஒाՉϕޑҢጄޣޑ ځύଫஒӃΕϕύǴԶќଫҢጄޣ߾ܫΕࡑύࡑǶ॥৻
໒௴ࡕǴᢀჸکϕ໔аϷϕکࡑ໔ޑ႖݈ӕਔܜ໒Ǵࡑ
ύޑҢጄޣΕϕࡕǴ൩ஒϕᆶࡑ໔ޑ႖݈ܫӣǶڙ၂ޣࡑӧᢀ ჸޑᢀჸਔ໔٩Ңጄޣޑ߄ԶۓǴҗჴᡍޣᒵҢጄޣ۶ԜޑϕݩǴ
Ңጄޣ۶ԜԖϕਔҔዸᒮՉीਔǹϕଶЗਔ߾ኩଶीਔǴҢጄޣ۶ ԜԖϕԶਔ໔ಕᑈԿ 1 ϩ 30 ࣾ܈ޣٿଫҢጄޣࡑӧϕϣޑᕴਔ໔ၲډ 29ϩដংǴջଶЗಕᑈਔ໔ǹ1 ϩដࡕǴϺޑᢀჸᏢಞ૽ግ൩ಖЗǶᢀჸѺ ର٣ҹಔޑҢጄޣϕۓကࣁǺيҽङඳࣁគৎޑҢጄޣటࠈќଫيҽङ ඳࣁᒡৎޑҢጄޣǴՠόхࡴគৎޑҢጄޣᒡৎޑҢጄޣᓸॹӧӦޑటϸ ᔐǴନߚϸᔐԋфωᆉǹᢀჸύ܄٣ҹಔޑҢጄޣϕۓက߾ࣁǺλ៓᠈ηѦޑ Ңጄޣჹ៓᠈ηϣޑҢጄޣՉޔௗޑ༘ᆪբǶᢀჸᏢಞ૽ግ่״ਔǴ॥৻
ᜢǴқՅό႖݈Ψӆӣᢀჸᆶϕϐ໔ǴฅࡕஒҢጄޣڗрǵӆڗ рڙ၂ޣǴӣচۚՐϐႵ᠈ϣǶᢀჸᏢಞ໘ࢤޑಃΒᆶಃΟϺ૽ግǴ܌Ԗჴᡍ ำׇᆶᢀჸᏢಞ໘ࢤಃϺ૽ግࢂֹӄ࣬ӕޑǶࣁΑዴᇡڙ၂ޣӧ૽ግಃϺа ϷനࡕϺޑѺର٣ҹύ܌ᢀჸډޑҢጄޣޑ߄όࢂԋࡕុޑᏫෳ၂໘ ࢤόӕಔڙ၂ޣϐϸᔈԖ܌ৡ౦ޑӢનǴӢԜךॺа૽ግಃϺаϷಃΟϺ܌