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標題: FXYD蛋白質之表現及調控廣鹽性硬骨魚(Tetraodon nigroviridis)滲透壓調節蛋白Na+, K+-ATPase之研究
FXYD proteins, the regulator of Na+, K+-ATPase of the euryhaline teleost, Tetraodon nigroviridis
作者: 王培任
Wang, Pei-Jen
關鍵字: FXYD
Tetraodon nigroviridis
出版社: 生命科學系所
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摘要: 本論文中主要的研究對象為墨綠凹鼻魨(Tetraodon nigroviridis)體內一種稱為FXYD的蛋白質(pFXYD),實驗目的是為了釐清pFXYD在廣鹽性硬骨魚滲透壓調節中所扮演的角色及其功用。 Na+/K+-ATPase(NKA)是一種特有的原生質膜酵素,能夠藉由主動運輸來排出鈉離子、並吸入鉀離子。NKA主要由α-subunit以及β-subunit兩種次單元組成。近年來在哺乳動物所發現的NKA γ-subunit擁有調控NKA活性的功能,NKA γ-subunit也被稱為FXYD2,是最早被發現的FXYD成員。先前在哺乳動物以及軟骨魚類的研究已經證實蛋白質FXYD是NKA的調節者,此發現引起我們想要探討當廣鹽性硬骨魚滲透壓調節器官中的NKA活性受到外界鹽度影響的時候,蛋白質FXYD的表現以及功用。 本研究中探討的三型墨綠凹鼻魨FXYD分別為pFXYD5、pFXYD8、pFXYD9,使用RT-PCR自滲透壓調節器官中得到DNA序列並且定序確認,之後將DNA序列轉譯成為蛋白質並且與其他脊椎動物的FXYD比對,接著畫出演化樹以確認蛋白質之間的親緣關係,三型pFXYD在鰓、腎、腸道等滲透壓調節器官中均有基因表現。pFXYD的mRNA表現量由real-time PCR測得,鰓上的pFXYD9以及腸道的pFXYD5、pFXYD8、pFXYD9都偵測到在淡水適應的河魨中有較高的表現量;相反地,腎臟的pFXYD9卻是在海水的環境中表現較多。實驗中依據pFXYD8和pFXYD9蛋白質的部分序列做為抗原來製作專一性抗體,pFXYD8和pFXYD9的抗體會在鰓以及腎中偵測到13 kDa的主要條帶(band),pFXYD9也會在腸道中被偵測到。此外,利用pFXYD5蛋白質的部分序列做為製作專一性抗體的抗原,在滲透壓調節器官鰓、腎、腸道中都會偵測到24 kDa的條帶。實驗結果發現,鰓上的pFXYD5和pFXYD9、腎臟的pFXYD5、腸道的pFXYD9在適應淡水環境時的表現量較高;另一方面,鰓的pFXYD8、腎臟的pFXYD8和pFXYD9、腸道的pFXYD5則是在海水中的表現量較高。蛋白質pFXYD在非洲爪蟾卵細胞(Xenopus oocyte)中過量表現的實驗證實了pFXYD5、pFXYD8以及pFXYD9均能抑制NKA活性。再以免疫螢光化學法偵測得鰓上的pFXYD9以及腎臟的pFXYD8皆與NKA分布在MR細胞上,共同免疫沉降法進一步證實了FXYD蛋白質與NKA之間的protein binding。將上述實驗結果與鰓、腎、腸道中的NKA活性相互比較並加以討論,本研究對於蛋白質pFXYD抑制NKA活性的特性,及其在廣鹽性硬骨魚滲透壓調節機制中所扮演的角色將提供一個更為全面性的論述。
In the present study, three FXYD protein members were studied in pufferfish termed pFXYD. The goals of the research were to elucidate the molecular and functional mechanisms of osmoregulation. Subsequently, the salinity-dependent responses of pFXYD proteins revealed their functional role in the euryhaline teleost (Tetraodon nigroviridis) when faced with an osmoregulatory challenge. The Na+/K+-ATPase (NKA) is a ubiquitous membrane-bound protein important for teleost osmoregulation. The enzyme is composed of two essential subunits; a catalytic α-subunit, and a glycosylated β-subunit responsible for membrane targeting of the enzyme. The smaller NKA γ-subunit, also known as FXYD2, is the first example of a small single transmembrane protein regulating NKA activity by interaction with the NKA α-subunit. Being the regulatory protein of NKA in mammals and elasmobranchs, it is intriguing to realize the expression and functions of FXYD protein in the euryhaline teleosts showing salinity-dependent changes of osmoregulatory organs NKA activity. The present study identified three cDNA sequences of pFXYDs confirmed by RT-PCR, including pFXYD5, pFXYD8, and pFXYD9. Amino acid sequences of pFXYD genes were deduced and the phylogenetical relationship of pFXYD proteins and other vertebrate FXYDs were analyzed. Pufferfish FXYD genes were expressed in the osmoregulatory organs of gill, kidney, and intestine of both freshwater (FW)- and seawater (SW)-acclimated euryhaline pufferfish. pFXYD9 showed differential expression across osmoregulatory tissues based on salinity acclimation. Based on real-time PCR, pFXYD9 was significantly higher in the gill and intestine of FW-acclimated pufferfish compared to SW-acclimated pufferfish. Conversely, renal mRNA expression of the pFXYD9 gene was higher in SW-acclimated fish when compared to FW-acclimated fish. On the other hand, pFXYD5 and pFXYD8 genes show a similar salinity acclimation dependent response in intestine only. Antibodies raised against partial amino acid sequences of the pFXYD8 and pFXYD9 proteins were detected in gill and kidney, however, in intestine, only pFXYD9 could be detected. On the other hand, an antibody raised against partial amino acid sequence of the pFXYD5 protein was applied in the immunoblots and a immunoreavtive band at 24 kDa was detected in three osmoregulatory organs. In branchial pFXYD5, branchial pFXYD9, renal pFXYD5, and intestine pFXYD9, the relative protein abundance was significantly higher in FW-acclimated group; but in branchial pFXYD8, renal pFXYD8, renal pFXYD9, and intestine pFXYD5 were significantly higher in SW-acclimated pufferfish. Function of pFXYD5, pFXYD8, and pFXYD9 all showed the ability to inhibit NKA activity by Xenopus oocyte experiments. Immunofluorescent staining of frozen sections provided direct evidence that branchial pFXYD9 and renal pFXYD8 were co-localized with NKA on MR cell. In addition, branchial pFXYD9 and renal pFXYD8 co-immunoprecipitated with NKA demonstrating their in vivo protein interaction. Comparisons of pFXYD mRNA and proteins expression with NKA activity were performed in gill, kidney, and intestine. These data provide evidence for the presences of pFXYD proteins, and their participation in osmoregulatory mechanisms by regulating NKA activity of the euryhaline teleost, Tetraodon nigroviridis.
其他識別: U0005-2008201009240500
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