臺灣博碩士論文加值系統

中文摘要
弓狀核的背中區是下視丘的局部多巴胺神經系統結節漏斗多巴胺（TIDA）神經系統所在。TIDA神經元活性有上午高下午低的日變週期現象，若破壞視叉上核（SCN），則TIDA神經元的週期現象便消失。此外，近年來本實驗室由側腦室注射血管活性腸胜（VIP）、腦下腺腺甘環化酶激活（PACAP）、蛙皮素（BB）等存在於SCN的神經胜，均可造成TIDA神經元的興奮。經由雙重免疫螢光染色法，可以在弓狀核的背中區，具酪胺酸羥化酶免疫活性的細胞旁，發現有VIP、PACAP及血管加壓素（AVP）免疫活性的神經末稍的分佈及並置，顯示確有VIP，PACAP及AVP之神經元投射至弓狀核。本論文之目的則是探討這些神經元源於何處。
論文中使用的動物模型，是採成熟的雌性大白鼠去卵巢後補充動情激素，此動物模型可有效模擬動情前期。在進行免疫染色前，有時會前處理秋水仙素，為了確定注射秋水仙素後，在不同的時間點犧牲動物，對TIDA神經元活性的週期變化是否有影響，以上述動物模型在補充動情激素後五天經由側腦室注射秋水仙素（48 g/rat），兩天後分別於1000、1500、1700快速斷頭犧牲動物，取其腦切片，以高效液相層析儀及電化學偵測系統檢測中突內多巴胺的代謝產物(DOPAC)濃度作為評估TIDA神經元活性的指標。實驗結果顯示在注射秋水仙素後，TIDA神經元的活性就不再呈現日變週期的改變，顯示神經元的活性表現，需要有軸突傳導的參與。本論文為顧及實驗的一致性，將實驗動物犧牲的時間點固定在上午進行。
為觀察有哪些神經核有神經纖維直接投射到弓狀核，則在補充動情素後第三天，以微量注射器將20 nl逆向神經追蹤劑fast blue直接注射到弓狀核，兩天後於側腦室注射秋水仙素，以阻斷軸突傳導，使欲觀察之神經胜能在細胞本體有較顯著的染色反應，再過兩天予以灌流犧牲，取前腦從前聯合至乳突體的腦切片進行螢光免疫染色，以螢光顯微鏡觀察有哪些神經核的VIP、PACAP、BB及AVP神經元直接投射到弓狀核。
實驗結果發現外側中膈區（lateral septum)、紋狀終底核(bed nucleus of the stria terminalis，BNST)、血管終板器(vascular organ of the lamina terminalis，OVLT)、中側視前區(medial preoptic area，MPOA)、穹窿下器(subfornical organ，SFO)、視丘室旁核(paraventricular nucleus of thalamus，PVT)、視上核(supraoptic nucleus，SON)、視叉上核(SCN)、下視丘室旁核(paraventricular nucleus of hypothalamus，PVN)、內側杏仁核(medial amygdala)及尾部弓狀核(caudal arcuate nucleus)的同側及對側神經核區域，均有神經元直接投射到弓狀核的注射部位。且位於MPO、SON、SCN、PVN的部份VIP神經元；位於MPO、SCN、PVN的部份PACAP神經元及位於SON、SCN、PVN的部份AVP神經元都有直接投射到弓狀核；唯獨BB沒有從上述的神經核區域中，被發現有直接投射到弓狀核。
由於含VIP、PACAP及AVP神經胜的神經元，多可由PVN直接投射到弓狀核，為進一步瞭解PVN神經元是否能直接與TIDA神經元形成突觸，故於動物去卵巢手術當天，同時將正向神經追蹤劑PHA-L直接注射到下視丘左側之PVN，第七天補充動情素，再過七天灌流犧牲，同樣取前腦切片進行雙重免疫組織化學染色法。結果除了同側（對側亦有少許）的Septum, BNST, OVLT, MPOA, PVT, SON, SCN, ME等區域可發現帶有PHA-L之神經纖維投射外，在弓狀核的背中區具酪胺酸羥化酶免疫活性的細胞旁，也發現許多有來自PVN的PHA-L免疫活性的神經末梢，顯示由PVN投射的神經纖維有與TIDA神經元直接形成突觸的可能。
綜合以上的結果顯示，弓狀核接受中樞許多神經核的投射，除了來自同側，也有許多來自對側神經核；同時在MPO、SON、SCN、PVN的部份含VIP、PACAP及AVP神經胜的神經元，可以直接投射到弓狀核，並可能藉此路徑調節TIDA神經元的活性。

英文摘要
A diurnal change of tuberoinfundibular dopaminergic (TIDA) neuronal activity is present in adult female rats regardless of their estrous stage. Lesion of the suprachiasmatic nuclei (SCN) invariably eliminates the rhythm. Our lab has shown that central administration of various neuropeptides, such as vasoactive intestinal peptide (VIP), pituitary adenylate cyclase-activating peptide (PACAP) or bombesin (BB), those originate in the SCN, results in potent stimulation of TIDA neuronal activity. Double fluorescent immunohistochemical staining revealed close apposition of VIP-, PACAP-, and vasopressin (AVP)-immunoreactive (ir) terminals with tyrosine hydroxylase (TH)-ir neurons within the dorsomedial arcuate nucleus (dmARN). We were then of interest to learn from where VIP-, PACAP- and AVP-containing neurons that project to the ARN originate.
Adult female Sprague-Dawley rats ovariectomized plus subcutaneous estrogen implants (OVX+E2) were used. Both retrograde and anterograde tracers were adopted and combined with immunohistochemical staining of various neuropeptides. Since colchicine, an axonal transport blocker, was used in the study to enhance the immunostaining of neuropeptides in the soma of neurons, it was also of interest to learn if colchicine can affect the diurnal change of TIDA neuronal activity. Intracerebroventricular injection of colchicine (48 g/rat) completely prevented the diurnal rhythm of TIDA neuronal activity 2 days later. The TIDA neuronal activity was determined by measuring 3,4-dihydroxyphenylacetic acid (DPOAC) levels in the median eminence using HPLC-ECD. This finding indicates that the rhythmic change of TIDA neuronal activity depends on neuronal inputs. Apparently, SCN is one of the sources. Although no diurnal change in TIDA neuronal activity was evident in colchicine-treated rats, all the rats used in this study were sacrificed by 1200 h to avoid possible other time-dependent variations.
The retrograde tracer, fast blue (FB), was microinjected into the ARN of OVX+E2 rats, and followed by colchicine 2 days later. After 2 more days, the rats were sacrificed by anesthesia and transcardially perfused with 4% paraformaldehyde. Their brains were removed, postfixed, cryosectioned, and mounted on slides. FB-labeled neurons observed with UV light under a fluorescent microscope were located in the septal region, bed nucleus of stria terminalis (BNST), organum vasculosa of lamina terminalis (OVLT), subfornical organ (SFO), medial preoptic area (MPOA), periventricular nucleus (PeVN), paraventricular nucleus of thalamus (PVT), SCN, supraoptic nucleus (SON), medial amygdala, paraventricular nucleus of hypothalamus (PVN) and caudal ARN. Most FB-labeled neurons were ipsilateral to the injection side; a few contralaterally labeled neurons were also present. The sections were further immunostained with VIP, PACAP, BB or AVP antiserum. Some FB-labeled neurons in the MPOA, SCN, PVN and SON were co-localized with VIP, PACAP or AVP. No co-localization of BB with FB-labeled neurons was observed in the brain regions studied.
Since PVN is one of the nuclei that contain the most FB-labeled neurons and also the double-labeled neurons, it was further studied with an anterograde tracer, Phaseolus vulgaris leucoagglutinin (PHA-L). PHA-L was injected directly into the PVN to examine the efferent projections of PVN and their possible connection with TIDA neurons in the ARN. Indeed, extensive PHA-L-ir terminals were found in the dmARN among other regions like lateral septum, BNST, OVLT, MPO, PeVN, PVT, SCN, SON, amygdala and PVN itself. Furthermore, many PHA-L-ir terminals exhibited direct contacts with TH-ir (presumed TIDA) neurons in the dmARN.
In conclusion: (1) ARN receives afferent inputs from various parts of the central nervous system, especially PVN, and (2) VIP-, PACAP- and AVP-containing neurons located in the MPOA, SCN, PVN and SON may play a role in regulating the TIDA neurons.

目錄
致謝………………………………………………………………………ii
中文摘要……………………………………………………………….. iii
英文摘要………………………………………………………………... v
第一章 緒論..……………………………………………………………1
第二章 材料與方法……………………………………………………18
第三章 實驗設計與結果………………………………………………26
第四章 討論……………………………………………………………30
圖表及說明……………………………………………………………..36
參考文獻………………………………………………………………..55

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