臺灣博碩士論文加值系統

本研究為能瞭解多巴胺在壓力源引發個體古典制約行為學習中的參與角色，採用一個與多巴胺相關的場地制約偏好行為作為研究工具，並利用一較溫和的禁錮壓力源作為非制約刺激與場地環境制約刺激進行配對制約。本研究假設內側前額葉皮質處的多巴胺參與在此壓力源引發場地制約行為學習中。實驗一針對單次禁錮壓力源的非制約刺激效果進行檢驗，分別檢測壓力源對個體的生理、情緒或是行為活動量的影響。實驗二利用「同時制約」或是「倒序制約」等兩種制約方式來進行單次禁錮壓力源引發場地制約偏好行為作業，並分別於制約程序的不同時間點施予多巴胺專屬受器拮抗劑，檢驗多巴胺在制約行為學習作業中的參與。實驗三在「同時制約」或是「倒序制約」兩種制約程序中的不同時間點，施予局部麻醉藥物二丁卡因暫時抑制內側前額葉皮質活動，以檢驗該區塊在單次禁錮壓力源引發場地制約偏好行為下的參與角色。實驗四為了解內側前額葉皮質處多巴胺在單次禁錮壓力源引發場地制約偏好行為中的角色，在「同時制約」程序的不同時間點施打多巴胺專屬受器拮抗劑至內側前額葉皮質區。
實驗結果發現：本研究所使用的單次30分鐘禁錮壓力源，確實可以引發實驗動物體內的壓力賀爾蒙糖皮質素大量增加、提高焦慮情緒或是降低自發性行為活動量。單次禁錮壓力源在「同時制約」或是「倒序制約」等兩種制約程序下，都能建立場地制約偏好行為。在禁錮壓力源操弄「之前」或「之後」，周邊施打多巴胺D1或是D2專屬受器拮抗劑，在「同時制約」或是「倒序制約」兩種制約程序中都會減抑禁錮壓力源建立場地制約偏好行為的效果。在「倒序制約」方式中，在「實驗動物接受完30分鐘壓力源操弄後被置入制約箱之前」才給予多巴胺拮抗劑，也會破壞後續的配對制約形成。在中樞內側前額葉皮質部分，在「同時制約」或是「倒序制約」兩種制約程序中，二丁卡因在壓力源與環境刺激配對「之前」給予才會抑制禁錮壓力源建立場地制約偏好行為的效果。在壓力源與環境刺激配對「之後」才抑制該處神經活動則不影響壓力源建立制約行為的效果。中樞內側前額葉皮質施予多巴胺D1或是D2專屬受器拮抗劑，也得到前述相同實驗結果。
本研究的實驗結果證明單次禁錮壓力源確實可以建立場地制約，為另類的古典制約行為。壓力源的操弄可引發多巴胺釋放量增加，及內側前額葉皮質處的多巴胺確實參與了此禁錮壓力源引發場地制約偏好行為。總結本研究結果顯示內側前額葉皮質處多巴胺在壓力下會對制約行為學習造成影響，並冀望此結果可以擴展對於內側前額葉皮質功能失能與心智疾患間關係的瞭解。在演化上，壓力對於人類或是其他族群的生存有其必要性。動物對於其環境中的危險或是威脅事件必須進行行為學習或因應，才能避免生命的損失。

To investigate the role of dopamine in stressor involved in classical conditioning, the present study used a dopamine-related task, conditioned place preference (CPP), as behavioral measurement. The mild restraint stressor was used and presumed to serve as the unconditioned stimulus to be paired with the contextual conditioned stimulus. The medial prefrontal cortex (mPFC) was hypothesized to be involved in this type of stressor induced place conditioning. Experiment 1 examined the effects of restraint stressor on physiological, emotional or locomotor tests. Experiment 2 investigated the involvement of dopamine in the stressor induced CPP, which conditioning procedures were manipulated by either simultaneous or backward form. The selective dopamine receptor antagonists were systemically administered in different time points during the conditioning procedures. Experiment 3 took lidocaine, a local anesthetic, to induce temporal deactivation of the mPFC. Lidocaine was infused in the mPFC at various time points, in either simultaneous or backward conditioning, to evaluate the involvement of the mPFC in stressor induced place conditioning. To further investigate the effects of dopamine receptors in the mPFC in the present type of CPP, the selective dopamine receptor antagonists were locally infused into the mPFC in simulutaneous conditioning procedure in Experiment 4.
The results showed that the manipulation of acute 30 min. restraint stressor increased the corticosterone, anxiety, but reduced the locomotor activities in rats. Consitent with previous work, this acute restraint stressor treatment given in either simultaneous or backward conditioning form significantly induced CPP. Systemic injection of dopamine D1 or D2 receptor antagonist given “before” or “after” the manipulation of restraint stressor, in either simultaneous or backward conditioning, attenuated the formation of stressor induced CPP. When these drugs were infused “right after the stressor manipulation and before the commencement of place conditioning” in the backward conditioning, the induction of CPP was also impaired. The attenuation of stressor formed place conditioning was showed when lidocaine was infused in the mPFC “before”, but not “after” the manipulation of restraint stressor. Such an attenuation effect was also seen when the selective D1 or D2 dopamine antagonist was infused in the mPFC.
The present study showed restraint stressor induced place conditioning as a novel type of classical conditioning. Consistent with the evidence showing that the manipulation of this stressor increases the release of dopamine, this study further verifed that the dopamine in the mPFC is involved in this restraint stressor induced CPP.

中文摘要…………………………………………………………………………..ⅰ
英文摘要…………………………………………………………………………..ⅲ
表目錄……………………………………………………………………..………ⅸ
圖目錄……………………………………………………………………………..ⅹ
第一章、緒論………………………………………………………………………1
第一節、多巴胺與酬賞典範…………………………………………………….…1
一、多巴胺神經傳導物質介紹……………………………….………………..…1
二、多巴胺與酬賞典範之建立：ahhedonia hypothesis…….…….…..…3
三、檢視多巴胺與酬賞典範…….……………………………………………..…4
（一）、重新檢視多巴胺與操作式制約行為的關係
（二）、多巴胺神經活動與酬賞物的時間鄰近性關係
（三）、多巴胺基因剔除小鼠與酬賞物關係之行為資料
（四）、酬賞物是否為一單一歷程？
（五）、重新檢視酬賞物概念
（六）、壓力源引發多巴胺釋放量增加現象
第二節、壓力源的影響………………………………………………………….11
一、壓力源對個體影響之理論………………………………………………….11
二、壓力源對個體行為學習的影響…………………………………………….13
（一）、壓力源特性對個體行為學習的影響
（二）、壓力源自身對個體行為學習的影響
第三節、壓力之相關神經機制……………………………………………………16
一、壓力賀爾蒙相關神經機制………………………………………………….16
（一）、下視丘-腦下垂體-腎上腺皮質的壓力賀爾蒙系統
（二）、壓力賀爾蒙與個體行為學習之關係
二、壓力與內側前額葉皮質……………………………………………………19
（一）、內側前額葉皮質神經解剖介紹
（二）、內側前額葉皮質與下視丘-腦下垂體-腎上腺皮質系統
（三）、內側前額葉皮質處多巴胺之相關功能探討
1、內側前額葉皮質與工作記憶
2、內側前額葉皮質與行為彈性
3、內側前額葉皮質與壓力因應
第四節、場地制約偏好行為………………………………………………………25
一、場地制約偏好行為…………………………………………………………25
（一）、自然酬賞物或藥物引發之場地制約偏好行為
（二）、非自然酬賞物或藥物引發之場地制約偏好行為
二、場地制約偏好行為作業的內容探討…………………………………………27
第五節、研究問題與實驗假設……………………………………………………29第二章、研究方法……………………………………………………………………31
第一節、受試者……………………………………………………………………31
第二節、工具………………………………………………………………………31
第三節、場地制約偏好行為的訓練步驟……………………………………….33
第四節、腦部埋管手術……………………………………………………………34
第五節、藥物………………………………………………………………………34
第六節、藥物顱內注射………………………………………………………….34
第七節、壓力賀爾蒙皮質醇酵素免疫分析之測定步驟…………………………35
第八節、組織切片…………………………………………………………………36
第九節、統計分析…………………………………………………………………36
第三章、實驗一：單次禁錮壓力源的非制約刺激效果檢驗……………..………38
第一節、實驗步驟…………………………………………………………………38
第二節、實驗結果…………………………………………………………………39
第三節、討論…………………………………………………………………….41
第四章、實驗二：多巴胺在單次禁錮壓力源引發場地制約偏好行為作業中的參與角色……………………………………………………………………………..43
第一節、實驗步驟…………………………………………………………………43
第二節、實驗結果…………………………………………………………………45
第三節、討論………………………………………………………………………49
第五章、實驗三：內側前額葉皮質在單次禁錮壓力源引發場地制約偏好行為下的參與角色………………………………………………………………………..54
第一節、實驗步驟……………………………………………………………...54
第二節、實驗結果…………………………………………………………………55
第三節、討論………………………………………………………………………60
第六章、實驗四：在單次禁錮壓力源引發場地制約偏好行為下，內側前額葉皮質處多巴胺的參與角色……………………………………………………..63
第一節、實驗步驟…………………………………………………………………63
第二節、實驗結果…………………………………………………………………63
第三節、討論…………………………………………………………………….65
第七章、綜合討論與結論…………………………………………………………67
第一節、壓力源建立場地制約偏好行為……………………………………….68
一、壓力源引發制約趨近行為………………………………………………….68
二、壓力源的非制約刺激效果………………………………………………….69
三、制約趨近行為的統計檢驗………………………………………………….74
第二節、壓力源的神經行為機制…………………………………………………75
一、多巴胺系統…………………………………………………………………75
1、多巴胺專屬受器拮抗劑對禁錮壓力源建立場地制約偏好行為的影響
2、多巴胺的角色功能再思考
二、壓力反應系統：內側前額葉皮質與壓力反應系統………………………79
第三節、結論…………………………………………………………………….82
參考文獻…………………………………………………………………………..84
附表………………………………………………………………………………105
附圖………………………………………………………………………………111

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