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系統識別號 U0026-2406201415262700
論文名稱(中文) 探討杏仁核與內嗅皮質功能性互動在古柯鹼學習與記憶之調節作用
論文名稱(英文) Functional Interaction between the Amygdala and Entorhinal Cortex in Modulating Cocaine Associated Learning and Memory
校院名稱 成功大學
系所名稱(中) 基礎醫學研究所
系所名稱(英) Institute of Basic Medical Sciences
學年度 102
學期 2
出版年 103
研究生(中文) 陳麗仙
研究生(英文) Li-Hsien Chen
學號 S58981191
學位類別 博士
語文別 英文
論文頁數 106頁
口試委員 指導教授-游一龍
口試委員-簡伯武
口試委員-徐嘉宏
口試委員-廖瑞銘
口試委員-吳豐森
口試委員-胡書榕
中文關鍵字 杏仁核  內嗅皮質  古柯鹼相關記憶  線索恐懼制約  情緒記憶 
英文關鍵字 amygdala  entorhinal cortex  cocaine associated memory and cue-induced fear conditionings 
學科別分類
中文摘要 中文摘要
在日常生活中附帶有情緒的事件,相較於平淡無奇的事件,通常會記得會比較鮮
明。神經解剖研究指出,杏仁核可藉由影響情緒喚起的程度來參與調控記憶的穩固化歷程。過去已知,杏仁核的功能是調控正面或負面情緒之記憶(包含藥物線索相關記憶與恐懼記憶)。本篇論文將進一步釐清杏仁核在調控古柯鹼線索相關記憶的穩固與再穩固化歷程中,以及杏仁核在線索恐懼記憶形成中,所扮演的角色。
在本論文的第一部分,本研究使用「古柯鹼制約場地偏好動物模型」來探討:杏仁核側底核區與內嗅皮質之間的互動在古柯鹼制約學習裡所扮演的角色。結果顯示,在古柯鹼制約學習之後,利用muscimol (一種GABAA受器的致效劑)加強杏仁核側底核區與內嗅皮質抑制性神經元的活性,會降低古柯鹼制約之場地偏好。此外,在古柯鹼制約場地偏好測試之後執行抑制或毀除杏仁核側底核區與內嗅皮質,也會減少之後古柯鹼制約場地偏好的表現。這些結果,皆指出杏仁核側底核區與內嗅皮質對於古柯鹼制約場地偏好的表現,是必要的。根據前述的發現,假設杏仁核側底核區與內嗅皮質互動進而調控古柯鹼制約場地偏好的再穩固歷程。因此,利用注射muscimol在同側單邊或對側單邊杏仁核側底核區與內嗅皮質,以檢驗兩核區之間的連結是否完整會有何影響。在古柯鹼制約場地偏好提取之後,於對側單邊杏仁核側底核區與內嗅皮質內注射muscimol,減少隨後古柯鹼制約場地偏好;然而,在古柯鹼制約場地偏好測試之後,於同側單邊杏仁核側底核區與內嗅皮質內注射muscimol,則不影響隨後古柯鹼制約場地偏好。最後,為了瞭解是否除了杏仁核側底核區與內嗅皮質外,仍有其他腦區參與古柯鹼制約場地偏好的形成。在提取古柯鹼制約場地偏好記憶之後,利用ibotenic acid毀除杏仁核側底核區與內嗅皮質,並不影響動物學習新的古柯鹼制約場地偏好。根據前述發現,假設應有其他腦區參與新的古柯鹼制約場地偏好的學習。為了檢驗這個假設,利用免疫組織方式檢測其他腦區Fos的表現。結果指出,在毀除杏仁核側底核區與內嗅皮質後,動物提取新的古柯鹼制約場地偏好,前額葉Fos蛋白質表現量有增加,而杏仁核中央核區Fos蛋白質表現量卻是減少的現象。
在此結論:杏仁核側底核區與內嗅皮質參與在古柯鹼記憶的表現。同側杏仁核側
底核區與內嗅皮質之間的神經互動參與調控古柯鹼制約場地偏好的再穩固歷程。在毀除杏仁核側底核區與內嗅皮質後,前額葉與杏仁核中央核區對新的古柯鹼制約場地偏好學習而言,是必要的。

第二部分,想要了解杏仁核在線索恐懼記憶的形成歷程是否有性別上的差異。在
本實驗將利用「線索恐懼制約小鼠動物模型」來檢驗創傷事件相關記憶的形成是否有性別差異。結果顯示,雌小鼠在線索制約僵直反應顯著地高過雄小鼠。利用皮下注射或是在杏仁核側核區注射雄性激素會減少性腺摘除的雄小鼠之線索制約僵直反應,然而雄性激素並不會影響性腺完好或性腺摘除雌小鼠的線索恐懼制約反應。因此,性別差異使雄性激素在雌雄小鼠線索恐懼記憶的表現有所不同。
英文摘要 ABSTRACT
Emotional arousal events tend to be remembered vividly than neutral ones. Neuroanatomical studies have suggested that amygdala is involved in memory consolidation by facilitating or dampening emotional arousal level. It is well known that the amygdala modulates positively and negative emotional memory, including drug-cues associated memory and fear memory. The present study aims to elucidate the role of amygdala in the consolidation, reconsolidation of cocaine associated memory and in the formation of cued-fear memory.
In part 1, we used cocaine-induced conditioned place preference to examine the role of functional interaction between the basolateral nucleus of amygdala (BLA) and entorhinal cortex (EC) in cocaine-conditioned memory. First, to elucidate whether BLA and EC are involved in the consolidation of cocaine-conditioned memory, we transiently inactivated BLA or EC with muscimol, a GABAA agonist, immediately after training. To further understand whether BLA and EC are critical for the retrieval of cocaine conditioned place preference, intra-BLA or -EC muscimol infusion prior to the CPP test decreased cocaine-induced CPP. Furthermore, Inactivation of BLA or EC immediately after the CPP test decreased subsequent performance in cocaine-induced CPP. These results, taken together, suggest that BLA and EC are necessary for cocaine-induced CPP performance. Based on our findings, we hypothesized that BLA in together with EC may modulate the reconsolidation of cocaine-induced CPP. To test the hypothesis, BLA and EC were functionally disconnected from each other by using unilateral muscimol infusion method. Importantly, ipsilateral intra-BLA and -EC muscimol infusion immediately after the CPP retrieval did not affect subsequent cocaine induced CPP. However, contralateral intra-BLA and -EC muscimol infusion immediately after the CPP retrieval drastically decreased subsequent cocaine-induced CPP. Finally, to examine whether is there a BLA- or EC- independent neural circuit for the formation of cocaine-induced CPP. After the retrieval of cocaine conditioned memory, the animals with BLA or EC lesion by ibotenic acid underwent new cocaine conditionings expressed the new cocaine-induced CPP. According to the finding, we supposed that there are complementary brain regions for the formation of new cocaine-induced CPP. For testing the hypothesis, the Fos immunostaining technique was used. The results revealed that the rats receiving new cocaine conditioning following BLA, EC, or BLA and EC lesion exhibited higher numbers of Fos-positive nuclei in the prefrontal cortex and less numbers of Fos-positive nuclei in the central nuclei of amygdala than sham control group.
We, hereby, conclude that BLA and EC are necessary for the storage of cocaine-induced CPP. Ipsilateral reciprocal neural activity between BLA and EC is involved in modulating the reconsolidation of cocaine-induced CPP. The PFC and are
essential for the formation of new cocaine-induced CPP after BLA and EC lesion.

In part 2, we want to understand whether there are sex differences in the activation of the amygdala during cue-induced fear conditionings processing. After cued fear
conditioning procedure, female mice exhibited a more robust cued fear memory than male mice. Systemic testosterone administrations and an intra-LA testosterone infusion
eliminated the CS-induced freezing in gonadectomized male mice, while testosterone administrations did not affect the CS-induced freezing in intact or OVX female mice.
Hence, these findings suggest that there is sex difference in the effect of testosterone on cued fear conditioning.
論文目次 TABLE OF CONTENTS
CHINESE ABSTRACT.......... II-III
ENGLISH ABSTRACT.......... IV-IV

Chapter 1 ............ 1
BACKGROUND ...........2-5

Chapter 2 ............7
Functional Interaction between Amygdala and Entorhinal Cortex in Modulating Cocaine Associated Learning and Memory
INTRODUCTION ............ 8-12
MATERIALS AND METHODS .........13-25
Animals ............13
Drug preparation and administration ....13
Stereotaxic Surgery and Cannula Implantation..14
Cocaine-induced CPP pretest, training, retest Pretest, training, test and retest protocol for cocaine-induced CPP experiments ............14
Procedures of Experiment 1.........16
Procedures of Experiment 2.........16
Procedures of Experiment 3.........19
Procedures of Experiment 4.........19
Procedures of Experiment 5.........20
Procedures of Experiment 6.........21
Procedures of Experiment 7.........22
Locomotor .............24
Histology .............24
Statistical Analysis...........25
RESULTS...............26
Experiment 1: Inhibition by infusing muscimol into bilateral BLA or EC immediately after cocaine-induced CPP training diminished cocaine-induced CPP....26
Experiment 2: The BLA and EC are associated with retrieval of cocaine-conditioned memory........27
Experiment 2.1: The current conditioning protocol rendered reliable Cocaine-CPP memory and performance.27
Experiment 2.2: Cocaine-CPP memory reactivation enhanced Fos expression in BLA and EC........28
Experiment 2.3: Cocaine-CPP memory reactivation did not affect Fos expression in the dorsal striatum....29
Experiment 3: Infusion of muscimol BLA or EC immediately before the test diminished the expression of cocaine-induced CPP.................30
Experiment 4: Bilateral infusion of muscimol into BLA and EC after the retrieval diminished cocaine-conditioned memory................30
Experiment 5: Contralateral intra-BLA and intra-EC muscimol infusion significantly attenuated cocaine-induced CPP.................31
Experiment 6: Role of the BLA and EC in Overtrained cocaine memory................32
Experiment 7: The BLA- or EC-independent neural circuit for cocaine associated learning.......34
Experiment 7.1: The animals with BLA, EC or both BLA and EC lesion underwent excessive alternation conditionings displayed new cocaine-induced CPP.......34
Experiment 7.2: mPFC may be a candidate complementary site after BLA or EC lesion for the formation of new cocaine-induced CPP...............35
DISCUSSION ..............37-41
REFERENCES ..............42-60
FIGURE LEGEND..............61-73

Chapter 3 ..............74
Sex Differences In Cued Fear Conditioning
INTRODUCTION.............75-77
MATERIALS AND METHODS..........78-83
Animals...............78
Drug preparation and administration.....78
Stereotaxic Surgery and Cannula Implantation..78
Determination of the Estrous Phase, Gonadectomy, and Gonadal Hormone Treatment.........79
Acoustic Stimulus (tone)- and Footshock-induced Freezing Responses...............81
Cued Fear Conditioning and Test.......82
Statistical Analysis..........83
RESULTS................84-26
Experiment 1: Tone- and Footshock-induced Freezing Response in the Male and Female Mice......84
Experiment 2: Cued Fear Conditioning in the Male and Female Mice...............85
Experiment 3: Gonadectomy and the Cued Fear Conditioning in the Male and Female Mice.........85
Experiment 4: Modulating Effects of E And 4-OHT on Cued Fear Conditioning in Ovariectomized Female and Male Mice.86
Experiment 5: Modulating Effects of T and Flu on Cued Fear Conditioning in Female and Gonadectomized Male Mice..87
DISCUSSION..............89-93
REFERENCES..............94-99
FIGURE LEGEND.............100-104

ABBREVIATIONS..............105-106 

FIGURES LIST
Chapter 1
BACKGROUND
Figure 1-1. Involvement of the amygdala in emotional memory..................5

Chapter 2
Functional Interaction between Amygdala and Entorhinal Cortex in Modulating Cocaine Associated Learning and Memory
Figure 2-1. Effects of intra-BLA and intra-EC muscimol infusions on the consolidation of cocaine-induced CPP.61
Figure 2-2. The development of a preference for the environment previously paired with cocaine....62
Figure 2-3. Schematic representation of coronal sections selected for quantification of Fos-positive nuclei..63
Figure 2-4. Numbers of Fos positive nuclei/0.36 mm2 in brain regions...............64
Figure 2-5. Effects of intra-BLA and intra-EC muscimol infusions on expression of cocaine-induced CPP. ..65
Figure 2-6. Effects of intra-BLA and intra-EC muscimol infusions on the reconsolidation of cocaine-induced CPP. ..................66
Figure 2-7. Effects of contralateral and ipsilateral intra-BLA and -EC muscimol infusion on the reconsolidation of cocaine-induced CPP.............67
Figure 2-8. Effects of intra-BLA and intra-EC ibotenic acid infusion after cocaine memory retrieval on subsequent cocaine conditionings...........68 
Figure 2-9. Effects of intra-BLA and intra-EC ibotenic acid infusion after cocaine memory retrieval on new cocaine conditionings...............69
Figure 2-10. Schematic representations of coronal sections for lesion in the basolateral nucleus of amygdala (BLA) and entorhinal cortex (EC)............70
Figure 2-11. Schematic representation of coronal sections selected for quantification of Fos-positive nuclei..71
Figure 2-12. Numbers of Fos positive nuclei/0.36 mm2 in brain regions...............72
Figure 2-13. Schematic diagram of the possible outputs and functions of these connections among the basolateral nucleus of amygdala and entorhinal cortex.......73

Chapter 3
Sex Differences In Cued Fear Conditioning
Figure 3-1. Tone- and footshock-induced freezing responses in male and female mice............100
Figure 3-2. Cued fear conditioning in male and female mice.................101
Figure 3-3. Gonadectomy and the Cued Fear Conditioning in the Male and Female Mice...........102
Figure 3-4. The modulating effects of Estradiol (E) and 4-hydroxytamoxifen (4-OHT) on the cued fear conditioning in ovariectomized (OVX) female and gonadectomized (TX) male mice.................103
Figure 3-5. The modulating effects of testosterone (T) and flutamide (F) on the cued fear conditioning in female and gonadectomized male mice...........104
參考文獻 Chapter 2
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Chapter 3
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