臺灣博碩士論文加值系統

硬骨魚生理易受環境因子 (溫度、鹽度、溶氧量、pH值、農藥和重金屬離子) 改變而影響其生理反應，其中鹽度改變造成的緊迫對魚體免疫反應的影響尚未清楚。因此，本實驗欲在急遽鹽度刺激下，利用MTT assay測定吳郭魚脾臟細胞添加mitogens (LPS或Con A)以探討其對吳郭魚脾臟細胞增生能力的影響。首先在無添加LPS或Con A刺激下，實驗結果脾臟免疫細胞增生能力在高滲透壓 (500 mOsmol/kg) 刺激1, 4, 8和24小時無顯著改變。分別給予不同濃度cortisol (20 nM和200 nM) 刺激後，發現給予200 nM cortisol刺激4小時後顯著抑制脾臟細胞增生。同時在高滲透壓 (500 mOsmol/kg) 和20 nM cortisol刺激4小時後，明顯抑制脾臟細胞增生。由此可知，高滲透壓刺激下，不會促使其脾臟細胞增生，而額外給予200 nM cortisol則明顯抑制脾臟細胞增生。另外，在添加LPS或Con A刺激的實驗中，在高滲透壓刺激1-8小時，脾臟細胞增生能力顯著增加；給予20 nM cortisol刺激1-4小時，脾臟細胞增生能力均明顯增加；而給予200 nM cortisol刺激4-8小時，脾臟細胞增生能力明顯增加；同時給予高滲透壓 (500 mOsmol/kg) 與20 nM cortisol刺激1-8小時後，脾臟細胞增生能力均明顯增加；若同時給予高滲透壓 (500 mOsmol/kg) 與200 nM cortisol刺激4小時，脾臟細胞增生能力才顯著增加。由上述可知，給予較低劑量之cortisol (20 nM) 抑制脾臟細胞增生能力較弱，因此在刺激1-4小時，發現仍能使脾臟細胞顯著增生；給予200 nM cortisol抑制脾臟細胞增生能力較強，但刺激4-8小時脾臟細胞顯著增生。在in vitro實驗中，cortisol 在不同濃度及作用時間，其調節脾臟細胞增生方式不同，推測cortisol可能經由刺激或改變其他細胞激素 (cytokines)，進而提高脾臟細胞增生。
本實驗將廣鹽性的莫三鼻克吳郭魚(Oreochromis mossambicus)由淡水轉移至25 ppt海水後不同時間，測定其plasma和脾臟lysozyme活性隨轉移時間持續顯著增加；鰓lysozyme活性則無顯著影響；頭腎lysozyme活性在轉移2小時內明顯增加。其次探討鹽度改變對吳郭魚脾臟中與stress反應有關之heat shock protein 70 (Hsp70) 和Hsp90、hemoglobin (Hb)、三種免疫相關性基因immunoglobulin M (IgM), major histocompatibility complex class Ι (MHC-I)、MHC classⅡ (MHC-Ⅱ)及基因表現的影響。將吳郭魚分別由淡水轉移至15、20、25、或35 ppt海水中2小時後，藉由半定量的RT-PCR/southern blotting方法分析Hsp70、Hsp90、Hb、IgM、MHC-Ⅰ及MHC-ⅡmRNA表現量，結果發現鹽度增加造成吳郭魚脾臟中Hsp70亦受不同鹽度的刺激後其表現量顯著增加，Hsp90在轉移至不同濃度海水2小時則無顯著改變。Hb、IgM、MHC-Ⅰ及MHC-Ⅱ mRNA表現量顯著高於長期淡水適應吳郭魚；而由此結果證實吳郭魚脾臟免疫相關基因的表現量在轉移至25 ppt海水2小時後有較顯著的影響。故將吳郭魚分別由淡水轉移至25 ppt海水1、2、4、8、16或 24小時，結果發現脾臟Hsp70在轉移4小時後表現量顯著增加。至於Hsp90表現量在轉移24小時和長期海水適應組才顯著增加。Hb mRNA表現量在轉移至25 ppt海水不同時間點均大於淡水組。IgM表現量在轉移至25 ppt 2小時達最大值 (2.22倍)，其餘時間點則均隨轉移時間而持續增加。MHC-Ι表現量隨轉移時間的增長其表現量逐漸增加。MHC-Ⅱ表現量隨轉移不同時間顯著增加。由此證明在急遽鹽度的刺激下，提高吳郭魚脾臟免疫基因IgM、MHC-Ⅰ和MHC-Ⅱ mRNA表達，以因應環境改變增強其免疫力。
利用2-D gel electrophoresis分離由淡水轉移至25 ppt海水4, 8和16小時後之吳郭魚脾臟蛋白質，並透過MALDI-TOF MS, ESI-MS/MS或MALDI-TOF/TOF MS鑑定出其中44種蛋白質 (33種蛋白質表現量為up-regulation，11種蛋白質表現量為down-regulation) 表現量受鹽度刺激而改變。其次以RT-PCR 分析insulin-like growth factor Ⅱ (IGF-Ⅱ)和ferritin middle subunit mRNA表現量，藉以確認蛋白質體分析結果之真偽及蛋白質與基因層次之相關性，結果IGF-ⅡmRNA level，在轉移至25 ppt海水不同時間，其表現量為up-regulation，且IGF-Ⅱ也在轉移1小時後極顯著的增加 (22-34倍)。在高滲透壓下，吳郭魚脾臟ferritin middle and light subunit mRNA和protein level均受鹽度刺激顯著提高，推測ferritin可能是一種適應osmotic stress蛋白質。由此可知2-D gel蛋白質體分析受改變之蛋白質其可信度高且與基因表達之改變一致。
高滲透壓刺激下，促使吳郭魚plasma和脾臟 lysozyme活性增加，亦增加脾臟免疫基因IgM、MHC-Ⅰ和MHC-Ⅱ mRNA表達。在高滲透壓下，cortisol亦抑制脾臟免疫細胞增生。從魚類適應高滲透壓調節機制角度得知，魚類在適應高滲透壓環境時，體內GH和cortisol分泌增加，藉以提升高滲透壓產生緊迫的適應能力，同時也增加魚體內的免疫能力，以利其達成適應高滲透壓的環境，使得魚得以存活。

The physiological regulations of teleosts are associated with the changes of environmental factors (temperature, salinity, oxygen, pH, chemical, heavy metals). When a euryhaline (broad salinity) teleost goes from hyposmotic (FW) to hyperosmotic (SW) environment, however the consequence of immunological response in tilapia remains unclear. This study was conducted to examine the lysozyme activity, functional genomic and proteomic changes in the spleen of tilapia (Oreochromis mossambicus) during acute salinity challenge. Isolated primary cultured splenocyte under pre-incubation with high osmolarity (500 mOsmol/kg) or cortisol (20 nM and 200 nM) for 1, 4, 8 and 24 h without or with mitogens (LPS or Con A) stimulation up to 96 h, the cell proliferation of tilapia splenocytes was measured by MTT assay. First, in the experiments without LPS or Con A, the data of the splenocytes proliferation were no significant difference when exposure to hyperosmotic (500 mOsmol/kg) medium. The proliferations of splenocytes pre- exposure to the 200 nM cortisol (in 300 mOsmol/kg) within 4 and 24 h were significantly inhibited. In addition, the proliferations of splenocytes pre-incubation with hyperosmotic added 20 nM cortisol medium within 4 h were significantly suppressed. On the other hand, the experiments with LPS or Con A, the proliferation of splenocytes when exposure to hyperosmotic (500 mOsmol/kg) medium for 1-8 h were significantly increased. Exposure with 20 nM cortisol for 1-4 h, the mitogen- stimulated proliferations of splenocytes were significantly increased while mitogen-stimulated proliferations of splenocytes were significantly increased in pre-incubation with 200 nM cortisol for 4 and 8 h. Moreover, pre-incubation with hyperosmotic added 20 nM cortisol for 1-8 h and with hyperosmotic added 200 nM cortisol within 4 h, the splenocytes proliferations were significantly elevated. This data imply that the profound effects of cortisol on the proliferation of splenocytes might mediate cytokines to enhance the stimulatory action of mitogens.
After exposure to various times of hyperosmotic (25 ppt SW) compared to fish kept in fresh water (FW), an increase significantly in plasma and spleen lysozyme activity were observed. No change was found in lysozyme activity of gill. In head kidney, lysozyme activity of SW was higher than those of FW within 2 h.
Under different salinities, Hsp70 mRNA of spleen was significant higher than that of FW. Hsp90 mRNA expression had no significant difference following directly transfer to SW. Tilapia acute transferred to various salinities (15, 20, 25 and 35 ppt SW) for 2 h, the increased expressions of Hb, IgM, MHC-I and MHC-ⅡmRNA in spleen were significantly correlated with salinity using RT-PCR/southern blotting. These results suggest that transfer to 25 ppt SW is significantly influenced the enhancement expression of spleen immune-related gene in tilapia. Furthermore, tilapia transferred to 25 ppt SW for 1, 2, 4, 8, 16 and 24 h, The results that after transfer to 25 ppt SW for 4 h, Hsp70 mRNA was significantly elevated in tilapia spleen. After transfer to 25 ppt SW for 24 h and long-term acclimation to SW, Hsp90 mRNA of tilapia spleen was significantly higher than that of FW. Furthermore, tilapia transferred to 25 ppt SW for 1, 2, 4, 8, 16 and 24 h, Hb mRNA level of spleen in 1 and 4-24 h of SW transfer tilapia were significantly higher than that of in FW. The peak (2.22 folds) expression of IgM within 2 h of SW transfer tilapia was significantly higher than that of FW acclimated fish, the remainders of time course maintained higher levels. MHC-I and MHC-Ⅱ mRNAs in spleen expressed significantly high with respect to the transfer time. These results suggest that the immune-related genes of tilapia spleen are induced under acute 25 ppt SW transfer.
Total proteins of spleen in tilapia transfer to 25 ppt SW for 4, 8 and 16 h were isolated and applied into 2-D gel electrophoresis with MALDI-TOF MS, ESI-MS/MS or MALDI-TOF/TOF MS to identify the profile and partial sequence of proteins. Compared with FW acclimated fish, 44 spots were differentially expressed: 33 spots belonged to be up-regulated and 11 spots were down-regulated. Two of identified proteins as IGF-Ⅱ and ferrintin middle subunit (FM) were further examined via mRNA expression to confirm the validation of proteomic analysis. The data showed that IGF-Ⅱ mRNA level was significantly increased in the spleen of tilapia after transferred from FW to 25 ppt SW for different time regimen. FM mRNA level in the spleen of tilapia followed SW transfer was associated with the increased levels of IGF-II mRNA. Moreover, Ferrintin light subunit (FL) protein (Western blotting) was significantly higher in the spleen of tilapia transfer to 25 ppt SW for 4 h than that of FW. The experimental data reveal that the evidence of IGF-II and FM mRNA expression provides a high correlate with RT-PCR. These results also suggest that IGF-II and FM may serve as a osmoregulated gene or protein to mediate the adaptation to hyperosmotic environment.
Taken the enhancements of plasma lysozyme activity, splenocyte proliferation, of spleen immune-related genes (IgM, MHC-I and MHC-Ⅱ) expression and of some identified proteins level (FM) altogether, tilapia can regulate the immunological responses and simultaneously enhance the salinity tolerance to adapt consequently the hyperosmotic conditions.

Index I
Content of tables III Content of figures IV
中文摘要 VII
Abstract IX
未來研究方向 XII
Introduction…………………………………………………………1
Osmoregulation………………………………………………………1
Immunomodulatory effects of GH and cortisol……………4
Lysozyme…………………………………………………………5
Heat shock protein 70 and 90 (Hsp70 and Hsp90)……6 Hemoglobin (Hb)……………………………………………7
Immunoglobulin M (IgM)…………………………………………8
Major histocompatibility complex class I and II (MHC-I and MHC-II)……9
Insulin-like growth factor I and II (IGF-I and IGF-II)…………………10
Ferritin………………………………………………………………12
Transferrin (TF)………………………………………………14
Specific aims………………………………………………………15
Experimental design………………………………………………16
Ⅰ. In vitro –MTT assay of tilapia splenocytes………16
Ⅱ. In vivo – measurement of lysozyme activity………17
Ⅲ. In vivo - mRNA expression………………………………18
Ⅳ. In vivo – proteomic analysis…………………………19
Materials and Methods………………………………………………...20
Preparation of fish splenocytes……………………………20
Splenocytes proliferation assay……………………………20
Experimental fish and osmotic stress………………22 Plasma collection…………22
Lysozyme activity………………………………………………22
Total RNA extraction and cDNA synthesis…………………23
PCR conditions…………………………………………………23
Southern blotting analyses…………………………………24
Spleen total protein extraction……………………………25
2-D electrophoresis……………………………………………25
In-gel digestion………………………………………………26
MALDI-TOF MS for protein identification…………………27
ESI-MS/MS of proteins…………………………………………27
SDS–PAGE gel electrophoresis………………………………27
Western blotting………………………………………………28
Statistical analysis…………………………………………28
Results…………………………………………………………………29
Discussion……………………………………………………………36
Conclusions…………………………………………………………46
References……………………………………………………………47
Content of tables
Table 1. List of specific primers used in this study……60
Table 2. Antibodies used in current investigations………61
Table 3. Lysozyme activity in tilapia plasma after the transfer from FW to 25 ppt SW……………………………………………………………………62
Table 4. List of protein identifications by MALDI-TOF MS..63
Table 5. List of protein identifications by MALDI-TOF MS…………………….67
Table 6. List of protein identifications by ESI-MS/MS…………………………..68
Table 7. List of protein identifications by ESI-MS/MS…………………………..69
Table 8. List of protein identifications by MALDI-TOF/TOF MS……………....70
Content of figures
Fig. 1. Effects of hyperosmotic medium (500 mOsmol/kg), cortisol (20 and 200 nM) and combinations of hyperosmotic and cortisol medium pre- incubation for 1, 4, 8 and 24 h on the proliferation of tilapia splenocytes without LPS or Con A stimulated………………………………………….71
Fig. 2. Effects of hyperosmotic (500 mOsmol/kg) pre-incubation for 1 (A), 4 (B), 8 (C) and 24 h (D) with LPS (10, 50 and 100 μg/ml) or Con A (1, 5 and 10 μg/ml)…………………………………………...……………………………72
Fig. 3. Effects of hyperosmotic medium (500 mOsmol/kg) pre-incubation for 1, 4, 8 and 24 h with LPS (10 μg/ml) or Con A (1 μg/ml)………………………74
Fig. 4. Effects of cortisol (20 and 200 nM) pre-incubation for 1 (A), 4 (B), 8 (C) and 24 h (D) with LPS (10, 50 and 100 μg/ml) or Con A (1, 5 and 10 μg/ml)…………………………...……………………………………………75
Fig. 5. Effects of cortisol (20 and 200 nM) pre-incubation for 1, 4, 8 and 24 h with LPS (10 μg/ml) (A) or Con A (1 μg/ml) (B)……………………...………...77
Fig. 6. Effects of hyperosmotic (500 mOsmol/kg) added cortisol (20 and 200 nM) pre-incubation for 1 (A), 4 (B), 8 (C) and 24 h (D) with LPS (10, 50 and 100 μg/ml) or Con A (1, 5 and 10 μg/ml)………………………………...…78
Fig. 7. Effects of osmolarity (500 mOsmol/kg) added cortisol (20 and 200 nM) pre-incubation for 1, 4, 8 and 24 h with LPS (10 μg/ml) (A) or Con A (1 μg/ml) (B)……………………...…………………………………………….80
Fig. 8. Effects of hyperosmotic (500 mOsmol/kg) and cortisol (20 and 200 nM) with LPS (10, 50 and 100 μg/ml) or Con A (1, 5 and 10 μg/ml) direct-incubation for 96 h………………………………...………………...81
Fig. 9. Effects of osmolarity (500 mOsmol/kg) and cortisol (20 and 200 nM) with LPS (50 μg/ml) or Con A (1 μg/ml) direct-incubation for 96 h……...…....83
Fig. 10. Effects of exposure to 25 ppt SW on plasma (A), gill (B), head kidney (pooled sample) (C) and spleen (D) lysozyme activity of tilapia……….84
Fig. 11. Hsp70 (505 bp) (A) and Hsp90 (751 bp) (B) mRNAs expression of spleen in tilapia exposed to different salinities (15, 20, 25 or 35 ppt SW) for 2 h…………………………………………………………………………….86
Fig. 12. Hb (334 bp) (A), IgM (255 bp) (B), MHC-Ⅰ(312 bp) (C) and MHC-Ⅱ(213 bp) (D) mRNAs expression of spleen in tilapia exposed to different salinities (15, 20, 25 or 35 ppt SW) for 2 h…………..………87
Fig. 13. Time-course expressions of Hsp70 (505 bp) (A) and Hsp90 (751 bp) (B) mRNAs abundance in spleen of tilapia exposed to 25 ppt SW………....88
Fig. 14. Time-course expressions of Hb (334 bp) (A) and IgM (255 bp) (B) mRNAs abundance in spleen of tilapia exposed to 25 ppt SW………....89
Fig. 15. Time-course expressions of MHC-Ι (312 bp) (A) and MHC-Ⅱ (213 bp) (B) mRNAs abundance in spleen of tilapia exposed to 25 ppt SW…….90
Fig. 16. Two-dimensional gel analysis of spleen in tilapia transferred from FW to 25 ppt SW for 4, 8 and 16 h………………………………………………91
Fig. 17. Two-dimensional gel analysis of spleen in tilapia transferred from FW to 25 ppt SW for 8 h………………………………………………………….94
Fig. 18. Tissue distributions of insulin-like growth factor II (IGF- II) and IGF-ⅠmRNA in tilapia exposed to 25 ppt SW for 4 h………………………….95
Fig. 19. Expressions of the insulin-like growth factor II (IGF- II) mRNA in the spleen of tilapia transfer to 25 ppt SW at various times……………..…96
Fig. 20. Two-dimensional gel analysis of spleen in tilapia transferred from FW to 25 ppt SW for 8 h………………………………………………………….97
Fig. 21. Two-dimensional gel analysis of spleen in tilapia transferred from FW to 25 ppt SW for 4 h………………………………………………………….98
Fig. 22. The alterations of ferritin middle subunit (FM) protein spot intensity in spleen of tilapia following 25 ppt SW transfer for various times (4, 8 or 16 h)…………………………..……………………………………………99
Fig. 23. Tissue distributions of ferritin middle subunit (FM), ferritin light subunit (FL) and transferrin (TF) mRNA in tilapia exposed to 25 ppt SW for 4 h……………………………………………………………….100
Fig. 24. Expressions of ferritin middle subunit (FM) (A) and ferritin light subunit (FL) (B) mRNA in the spleen of tilapia transfer to 25 ppt SW at various times………………………………………………………………………101
Fig. 25. Ferritin light subunit (FL) protein levels in the spleen of tilapia transfer to 25 ppt SW at various times (4, 8 or 16 h)……………………………102

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