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|標題:||Functional annotation of proteomic analysis from the hypothalamus of Leghorn layers by thyroxine induction|
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|摘要:||動物藉由感受到日照長短來調整生殖、遷徙、冬眠和換羽等生理行為。此種隨季節不同而有行為改變的現象稱作光週期性(photoperiodism)。光接受器感受光刺激，此訊息會傳至中心樞生理時鐘，經綜合與轉譯這些訊號後，以神經內分泌反應輸出，以激活相關反應與功能諸如生殖生理與行為。甲狀腺素(thyroid hormone)除了主要調控能量代謝外，還能媒介光照反應來生殖生理行為。下視丘中基部(mediobasal hypothalamus, MBH)是調控鳥類日夜節律(circadian rhythm)的中心位置，其中type 2 iodothyronine deiodinase (Dio2)會將thyroxine (T4) 由prohormone狀態轉化成有生物活性的三碘甲狀腺酪胺酸(3, 5, 3'–triiodothyronine, T3)，進而刺激下視丘分泌促性腺刺激素釋放激素(gonadotropin-releasing hormone, GnRH)的神經末梢擴展接近腦下垂體門脈系統以利GnRH由下視丘釋放，GnRH再促進腦下垂體-性腺軸線(pituitary-gonad)。c-Fos在被激活神經元去極化後大量表現，其蛋白質產物 c-Fos 被當作是辨認神經元細胞活化的生物標記。因此本研究以蛋白質體學方法探討T3對禽類下視丘神經元與GnRH的分泌之影響路徑。試驗使用50週齡來航雞，於夏季光照進入黑暗時期(傍晚六點)翼靜脈注射T3 (5 μg/kg body weight)或生理食鹽水(1 mL/kg body weight)分別作為處理組和控制組，4小時後犧牲取其下視丘供蛋白質體分析。結果顯示: 在所有定量的420個蛋白質點中有14個蛋白質點有顯著差異(P < 0.05)，其中有六個蛋白質點表現量上升，另外八個蛋白質點表現量下降，這些有差異的蛋白質多與調控能量代謝相關路徑及細胞骨架的形成有關，以西方吸漬法進一步確認丙酮酸激酶(pyruvate kinase)蛋白質表現，的確於T3處理後顯著下降(P<0.05)。免疫組織螢光染色法分析發現，T3處理下，c-Fos在的第三腦室周圍神經元細胞大量表現，且c-Fos和GnRH兩者具有共同的神經元表現位置，西方吸漬法進一步確認 T3處理後下視丘中c-Fos的表現顯著高於控制組(P < 0.05)。過去研究顯示: 由白天進入晚上時由周邊施打甲狀腺素後，蛋雞下視丘能量感測和調節相關基因(UCP, SIRT, LKB, PGC-1α)表現脈動提早，以產生能量應付活動作狀態，而晝夜節律基因CLOK、BMAL1在白天時表現量有較顯著的波動，綜何過去與本研究結果可推測周邊循環T3可藉由直接影響下視丘或經影響身體周邊組織能量狀態後，以能量缺乏訊號刺激下視丘，活化下視丘神經元來調控GnRH表現。|
Animals can adjust their physical, physiological, and behavioral responses such as reproduction, migration, hibernation, and molting, in response to changing lengths of sun light. Behavioral adjustment in adaption to seasonal changes is called photoperiodism. Photoreceptors detect stimulation of light. A central clock or timer translates the signals. Then the outputs of neuroendocrine information activate behaviors and ally physiological responses such as reproductive functions. In addition to the effect on energy homeostasis, thyroid hormone can also regulate reproduction. Mediobasal hypothalamus is an important center controlling photoperiodic rhythm in avian species. The level of type 2 iodothyronine deiodinase (Dio2) in the mediobasal hypothalamus increases under the long-day photoperiod to catalyze thyroxine (T4) to its active form, 3,5,3'–triiodothyronine (T3), which in turn stimulates gonadotropin-releasing hormone (GnRH) nerve terminals located in close contact with the basal lamina for GnRH release, which then go through the hypophysial portal system to activate the pituitary-gonad axis. The c-Fos is highly expressed after neuron depolarization, and its protein product, c-Fos, tends to be used as a biomarker of neuron activation. Therefore, the aim of this study was to delineate the possible mechanisms how circulating T3 exerts effects on the hypothalamic neurons in relation to the secretion of GnRH by proteomic approaches. Leghorn hens at 50 weeks of age were given T3 (5μg/kg) or saline (1 mL/kg body weight) by intravenous injection before dark (at 6 o'clock, PM). The hypothalamuses were collected for proteomic analysis 4 hours after treatment. Results showed that 14 out of 420 quantified protein spots differed significantly by T3 induction (P < 0.05). Among the differentially expressed proteins, 6 and 8 spots were upregulated and downregulated, respectively. Most of the differentially expressed proteins are involved in metabolic pathways of energy homeostasis and formation and mobilization of cytoskeletons. Western blotting analysis confirmed the downregulation of pyruvate kinase expression by T3 induction as observed in proteomic analysis (P<0.05). Immunohistochemical studies further showed that T3 treatment promoted c-Fos expression, which were highly located in the neurons along the third ventricle, and were co-localized with GnRH-releasing neurons. Significant activation of c-Fos in the hypothalamus by T3 induction was further confirmed by Western blotting analysis (P < 0.05). Past studies suggested that T3 stimulation in hens mimics the light cue, which in turn drives the hypothalamus to reply an early ignition and fluctuation of energy-sensing and regulatory gene expressions including liver kinase B1 (LKB1), uncoupling protein (UCP), NAD-dependent deacetylase sirtuin-1 (SIRT1), and peroxisome proliferator-activated receptor-γ coactivator-1 alpha (PGC-1α). T3 treatment also induced a more fluctuating pattern of CLOCK and BMAL1 expression during the day. In combination with the results in the current study, it was concluded that circulating T3 may affect hypothalamic activation and following GnRH secretion pulse by directly acting on the hypothalamus or via a feedback loop as a low energy status from peripheral tissues.
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