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研究生： 余禮儒
Yu, Li-Lu
論文名稱： 廣鹽性青鱂魚仔魚體表離子細胞排放鉀離子之機制
Potassium Secretion by Ionocytes in the Skin of Medaka Larvae
指導教授： 林豊益
Lin, Li-Yih
學位類別： 碩士
Master
系所名稱： 生命科學系
Department of Life Science
論文出版年： 2015
畢業學年度： 103
語文別： 英文
論文頁數： 59
中文關鍵詞： 魚類 、離子調控 、離子細胞 、鉀離子 、青鱂魚
英文關鍵詞： fish, ion regulation, ionocyte, potassium, medaka
論文種類： 學術論文
相關次數： 點閱：198  下載：0 
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    • 魚類鰓與皮膚上的離子細胞是維持離子與滲透壓恆定的主要細胞。離子細胞參與鈉、氯、鈣離子的調節機制已經有許多研究，但是脊椎動物身體內重要的陽離子¬鉀離子的調節機制卻不甚明確。近期研究發現，斑馬魚離子細胞中有一群細胞的頂膜會表現鉀離子通道(ROMK)，而底側膜有鈉鉀幫浦(NKA)。在吳郭魚(Mozambique tilapia)的研究也發現類似的細胞，並且推論這種細胞是參與鉀離子分泌的離子細胞。然而，這些研究仍然缺少直接的證據去證實鉀離子分泌細胞的功能。本研究使用廣鹽性青鱂魚(Oryzias latipes)作為模式物種，利用掃描式離子選擇電極技術(SIET)測量仔魚皮膚上離子細胞鉀離子流，證實有一群離子細胞會分泌鉀離子。高鉀淡水與海水環境的馴養後會促進鉀離子的分泌。而且鉀離子的分泌會受到ROMK與NKCC抑制劑的抑制。利用原位雜交與細胞免疫組織染色也發現ROMKa確實會表現在離子細胞中。定量PCR分析魚鰓上mRNA的變化，發現高鉀環境馴養能夠增加ROMKa與NKCC1a的表現量。本研究提供直接的證據支持離子細胞分泌鉀離子的生理功能，並提出ROMKa與NKCC1參與鉀分泌的機制。

    The ionocyte in gills and skin of fish plays a critical role in ionic and osmotic regulation. Molecular mechanisms of sodium, chloride and calcium regulation in the ionocytes have been well investigated. However, the regulatory mechanism of potassium, which is a major monovalent cation in fish, is still unclear. In recent studies, a sub-group of ionocyte, which expresses a renal outer medullar potassium channel (ROMK) in the apical membrane and Na+/K+-ATPase (NKA) in the basolateral membrane was identified in zebrafish and Mozambique tilapia, suggesting that those ionocytes are involved in potassium secretion. However, the potassium secreting function of ionocyte is not fully understood. In this study, we investigated the mechanism of potassium secretion by using medaka (Oryzias latipes) as an animal model. With a scanning ion-selective electrode technique (SIET), potassium secretion was detected at the apical surface of a group of ionocytes in the skin of medaka larvae. The potassium secretion was enhanced after high potassium water (HK) or seawater acclimation, and was inhibited by ROMK and Na+-K+-Cl- cotransporter (NKCC) inhibitors. In situ hybridization and immunohistochemistry showed that kcnj1a and NKCC1a were expressed in the skin ionocytes. Quantitative PCR showed that mRNA levels of kcnj1a and nkcc1a were induced by HK acclimation. Taken together, this study provides physiological and molecular evidence to show that ROMKa and NKCC1 are involved in the potassium secretion of ionocytes.

    摘要 5 Abstract 6 Introduction 7 The importance of potassium in vertebrate animals 7 Potassium regulation in terrestrial animals 7 Na+-K+-Cl- cotransporter in renal potassium reabsorption 8 Renal outer medullar potassium channel in renal potassium secretion 9 Potassium disorder 10 Ion-regulation in fish 10 Ionocytes in fish skin and gill 11 Renal outer medullar potassium channel (ROMK) in fish gills 12 Na+-K+-Cl- cotransporter in fish gills 13 Scanning ion-selective electrode technique (SIET) and animal model 13 Purposes and Hypotheses 15 Experimental Designs 16 Experiment 1. Whole-body K+ and Na+ contents in different stages of medaka larvae acclimated to KF, NW or HK 17 Experiment 2. K+ gradient at skin surface of medaka larvae 17 Experiment 3. K+ gradient at the yolk-sac surface in larvae acclimated to KF, NW and HK 17 Experiment 4. K+ fluxes at individual ionocytes and keratinocytes in the yolk-sac skin of larvae 17 Experiment 5. Effect of short time high ammonia on K+ and NH4+ fluxes in ionocytes and keratinocytes of larvae 18 Experiment 6. Effects of ROMK inhibitor (VU591) on K+ transport 18 Experiment 7. Effects of NKCC inhibitor (bumetanide) on K+ transport 18 Experiment 8. Quantitative real time-PCR analysis of gene expression in gills of medaka acclimated to NW or HK 18 Experiment 9. In situ hybridization of kcnj1a and immunohistochemistry of NKA in ionocytes of larvae 19 Materials and Methods 20 Experimental animals 20 Acclimation experiments 20 Measurement of whole-body K+ or Na+ contents in medaka larvae 21 Scanning ion-selective electrode technique (SIET) 21 Measurement of surface K+ gradients 22 Measurement of cellular K+ fluxes for specific cells 23 Treatments of VU591 and Bumetanide 24 Preparation of RNA 24 Quantitative real-time polymerase chain reaction (qRT-PCR) analysis 25 RNA probe synthesis 26 In situ hybridization and immunohistochemistry (IHC) 27 Statistical analysis 28 Results 29 Experiment 1. Whole-body K+ and Na+ contents in different stages of medaka larvae acclimated to KF, NW or HK 29 Experiment 2. K+ gradient at skin surface of medaka larvae 29 Experiment 3. K+ gradient at the yolk-sac surface in larvae acclimated to KF, NW and HK 30 Experiment 4. K+ fluxes at individual ionocytes and keratinocytes in the yolk-sac skin of larvae 30 Experiment 5. Effect of short time high ammonia on K+ and NH4+ fluxes in ionocytes and keratinocytes of larvae 30 Experiment 6. Effects of ROMK inhibitor (VU591) on K+ transport 31 Experiment 7. Effects of NKCC inhibitor (bumetanide) on K+ transport 31 Experiment 8. Quantitative real time-PCR analysis of gene expression in gills of medaka acclimated to NW or HK 32 Experiment 9. In situ hybridization of kcnjk1a and immunohistochemistry of NKA in ionocytes of larvae 32 Discussion 33 K+ content of larva 33 K+ secretion by larval skin 33 K+ secretion in larvae acclimated to different K+ levels 34 ROMK inhibitors 35 Effect of NH4+ on K+ secretion 36 Role of ROMK in fish ionocyte 36 ROMK in subtypes of ionocytes 37 NKCC role in medaka ionocyte 38 Conclusion 39 References 40 Figures 48 Fig. 1. Whole-body K+ (A) or Na+ (B) contents in different stages of medaka larvae acclimated to potassium-free water (KF), normal-fresh water (NW) or high-potassium water (HK). 48 Fig. 2. K+ gradient at skin surface of 7 dpf medaka larvae. 49 Fig. 3. K+ gradient at yolk-sac surface of 7 dpf medaka larvae acclimated to potassium-free water (KF), normal-fresh water (NW) or high-potassium water (HK). 50 Fig. 4. K+ flux at ionocytes (IC) or keratinocytes (KC) of 7 dpf medaka larvae acclimated to normal-fresh water (NW) or 15 ‰ and 30 ‰ salinity seawater (15 SW, 30 SW). 51 Fig. 5. Short time high ammonia effect of K+ (A) or NH4+ (B) fluxes at ionocytes (IC) or keratinocytes (KC) of 7 dpf medaka larvae acclimated to normal-fresh water (NW). 53 Fig. 6. Effects of ROMK inhibitor (VU591) on K+ gradient at the yolk-sac surface (A) or ionocytes (IC, B) and keratinocytes (KC, B) of 7 dpf medaka larvae acclimated to normal-fresh water (NW). 54 Fig. 7. Effects of NKCC inhibitor (bumetanide) on K+ gradient at the yolk-sac surface (A) or ionocytes (IC, B) and keratinocytes (KC, B) of 7 dpf medaka larvae acclimated to normal-fresh water (NW). 55 Fig. 8. Quantitative real-time PCR analysis of kcnj1a, 1b (A) and nkcc1a, 1b (B) relative mRNA expression in gills of medaka acclimated to normal-fresh water (NW) or high-potassium water (HK). 56 Fig. 9. in situ hybridization and immunohistochemistry of 7 dpf medaka larvae acclimated to normal-fresh water (NW). 57 Fig. 10. The putative model of K+ secretion by FW-type ionocytes of medaka. 59

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