結論

一、本研究已成功選殖到白色來亨雞之端粒反轉錄酶 (telomerase reverse transcriptase, TERT) 基 因 並 進 行 DNA 序 列 分 析 比 對，期能供基因轉殖並發展延長家禽體細胞體外培養的技術平 台。

二、 本研究已完成 畜產試驗 所應用體 細胞核 轉置 (somatic cell

nuclear transfer, SCNT) 技術生產的複製荷蘭種乳牛與阿爾拜 因乳山羊及其後代動物在不同特定年齡的端粒 (telomere) 長 度，發現複製牛與正常同齡動物並無顯著差異存在，本研究證 明應用 SCNT 技術生產的複製牛，其調控端粒長度的相關基 因可以順利的進行再程式化，使端粒維持一定的長度水準，並 沒有因為使用成體之耳朵成纖維細胞做為供核細胞而導致端粒 短化的老化現象。但複製羊則顯著（p＜0.05）較正常同齡對照 羊為短。本研究顯示，應用 SCNT 技術生產的複製羊其端粒長 度較正常對照為短，隨著月齡的增加，端粒的長度顯著較對照 羊為短。應用複製技術生產的乳牛及乳羊，可能因為動物種別 不同，其端粒長度在經過 NT 操作後，會因為胚在發育過程其 染色體再程式化程度之不同，使端粒長度回復正常或較正常動 物為短。

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表 1. 白色來亨雞全長 TERT 之 DNA 序列進行 BLAST 比對後依 序列相似度排序的結果

Table 1. Ranking of identity of White Leghorn chicken TERT related genes according to BLAST analysis.

Access no. Description Max score

Gallus gallus telomerase reverse transcriptase (TERT), mRNA >gb|AY626231.1| Gallus gallus telomerase reverse transcriptase (TERT) mRNA, complete cds

8715 8715 100% 99%

AY502592.1 Gallus gallus gallus telomerase reverse

transcriptase (TERT) mRNA, complete cds 8102 8102 93% 99%

DQ681294.1

Coturnix japonica telomerase reverse transcriptase (TERT) mRNA, complete cds, alternatively spliced

6806 6806 93% 94%

EF552224.1 Gallus gallus clone chTERTAC-GSC

telomerase catalytic subunit mRNA, partial cds 3349 3349 38% 100%

DQ681293.1

Anas platyrhynchos telomerase reverse transcriptase (TERT) mRNA, partial cds, alternatively spliced

2143 2143 47% 84%

BX934708.1 Gallus gallus finished cDNA, clone

ChEST116f8 2006 2006 23% 99%

BX934910.2 Gallus gallus finished cDNA, clone

ChEST151a12 2002 2002 23% 99%

EF552225.1

Gallus gallus clone chTERTAC-MDCC-MSB1 nonfunctional telomerase catalytic subunit mRNA, partial sequence

1236 2843 32% 100%

DQ681312.1

Gallus gallus telomerase reverse transcriptase isoform K (TERT) mRNA, partial cds, alternatively spliced

968 1329 15% 100%

DQ681309.1

Coturnix japonica telomerase reverse transcriptase isoform qK3 (TERT) mRNA, partial cds, alternatively spliced

915 915 13% 93%

AY505016.1 Gallus gallus gallus telomerase reverse

transcriptase (TERT) gene, 3' UTR 911 911 10% 99%

DQ681298.1

Gallus gallus telomerase reverse transcriptase isoform J (TERT) mRNA, partial cds, alternatively spliced

710 1241 14% 100%

DQ256207.1

Gallus gallus telomerase reverse transcriptase variant 1 (TERT) mRNA, exons 2 through 4 and partial cds, alternatively spliced

697 777 8% 100%

DQ681297.1

Gallus gallus telomerase reverse transcriptase isoform O (TERT) mRNA, partial cds, alternatively spliced

691 1174 13% 100%

DQ681313.1

Gallus gallus telomerase reverse transcriptase isoform E2 (TERT) mRNA, partial cds, alternatively spliced

682 842 9% 100%

DQ256206.1

Gallus gallus telomerase reverse transcriptase variant 1 (TERT) mRNA, exons 2 through 4 and partial cds, alternatively spliced

680 744 8% 100%

DQ256209.1

Gallus gallus telomerase reverse transcriptase variant 1 (TERT) mRNA, exons 3, 4 and partial cds, alternatively spliced

678 678 7% 100%

表 2. 與白色來亨雞 TERT 蛋白質具有最高相似性的 10 種蛋白質序列 Table 2. Top ten protein sequences which share the highest amino acid

similarity with the White Leghorn chicken TERT.

Access no. Description Similarity

ref|NP_001026178.1| telomerase reverse transcriptase [Gallus

gallus]

1343/1346

(99%) gb|AAS75793.1| telomerase reverse transcriptase [Gallus

gallus gallus]

1340/1346

(99%) gb|ABG75863.1| telomerase reverse transcriptase

[Coturnix japonica]

1263/1347 (93%) ref|XP_001369432.1| PREDICTED: similar to telomerase

reverse transcriptase [Monodelphis

domestica]

672/1347 (49%) gb|ABQ15326.1| telomerase catalytic subunit [Gallus

gallus]

604/604 (100%) gb|ABG75862.1| telomerase reverse transcriptase [Anas

platyrhynchos]

485/613

(79%) ref|NP_001079102.1| telomerase reverse transcriptase [Xenopus

laevis]

486/865 (56%) ref|NP_937983.2| telomerase reverse transcriptase isoform

1 [Homo sapiens]

456/860 (53%) gb|AAC40212.1| telomerase catalytic subunit [synthetic

construct] 455/860

(52%) gb|AAC51724.1| telomerase catalytic subunit [Homo

sapiens]

455/860

(52%)

表 3. 複製牛及其後代與對照牛之基本資料

Table 3. The background information of cloned cattle, offspring, and age-match control animals.

動物個體 性別 出生日期 採血進行端粒

No. 468 Female 2006.01.30 3.0 No. 393 Male 2005.09.18 7.0

No. 444 Female 2005.10.08 6.0 No. 312 Female 2004.02.10 24.0

＊

採血日期為 2006 年 04 月 18 日

NB：採血時剛出生 (new born)

表 4. 複製羊及其後代之基本資料

Table 4. The background information of cloned dairy goats and their offspring.

動物個體 性別 出生日期 採血進行端粒

分析時年齡

^＊

SCNT 乳羊 月齡

寶鈺 Female 2004.03.10 25 寶祥 Female 2005.04.25 50 HAS

^＊＊

基因轉殖複製羊 Female 2005.08.07 8 SCNT 乳羊後代

寶貝 Male 2005.04.25 12

寶鈺之女 1 號 Female 2006.03.04 17 寶鈺之女 2 號 Female 2006.03.04 17

寶祥之子 Male 2005.02.27 14

寶祥之女 Female 2005.02.27 14

＊

採血日期為 2006 年 04 月 18 日

＊＊

人類血清白蛋白 (human serum albumin, HSA)

表 5. 複製牛及其後代與同年齡非複製牛之端粒長度

Table 5. Telomere length of cloned cattle, offspring, and age-match control animals.

動物別 數量

(N)

端粒長度 (kb) Mean ± SD 複製牛^＊ 4 17.52 ± 0.41 複製牛同齡對照牛 4 17.66 ± 0.32 複製牛後代 4 17.67 ± 0.32 複製牛後代同齡對照牛 5 18.69 ± 0.50

供核體細胞 (HE 921) 1 17.30

複製牛^＊＊ 4 17.48 ± 0.31 複製牛同齡對照牛 6 17.80 ± 1.57

供核體細胞 (HE 921) 1 17.07

＊

採血日期為 2006 年 04 月 18 日

＊＊

採血日期為 2007 年 01 月 08 日

表 6. 複製羊及其後代與同年齡非複製羊之端粒長度

Table 6. Telomere lengthes of cloned dairy goats, offspring, and age-matched control animals.

複製羊 月齡 (months) 端粒長度 (kb) 同月齡對照羊 端粒長度 (kb)

(N=3)

1.(

Bao Siang

) 50 16.68 16.92±0.73

2.(Paoyu) 22 17.05 17.04±0.62

3.(HAS tg) 6 14.32

^a

17.37±0.32

^b

1.(

Bao Siang

) 62 12.26

^a

16.08±0.86

^b

2.(Paoyu) 34 13.50

^a

15.72±0.61

^b

3.(HAS tg) 18 10.39

^a

16.32±0.61

^b

複製羊後代 (N=5) 12-17 16.07 ± 0.32 16.43 ± 0.36

供核體細胞 (GE 1244, P2)

14.86

供核體細胞 (GE 09181, P8)

14.20

Letters a and b in same row indicate significantly different ( p ＜ 0.05)

圖 1. 應用顯微抽取技術自白色來亨雞蛋取得原腸期雞胚樣品

Fig. 1. Collection of gastrula stage chicken embryos from White Leghorn chicken by micromanipulation.

2150~3358 3057~4764 1001~2203

Promoter

44~1052

圖 2. 參考網路資料庫 AY626231 之序列進行 PCR 擴增自 5’至 3’

的部分基因片段產物

Fig. 2. The partial PCR fragments located from 5’ through 3’ were amplified according to the Gen Bank no. AY626231 sequence.

M E PGCs Nc E PGCs

500 bp 1000 bp

508 bp TERT 283 bp Gapdh

圖 3. 自白色來亨雞胚 (E) 與始基生殖細胞 (PGC) 樣品中擴增家禽 端粒酶基因及陽性對照GAPDH (glyceraldehyde-3-phosphate dehydrogenase) 的部分基因片段產物。M為 100 bp 標記。Nc 為陰性對照組。

Fig. 3. Electrophoresis analysis of GAPDH and TERT PCR products amplified from gastrula stage embryos (E) in White Leghorn chicken. M: 100 bp ladder marker. Nc: negative control.

- 3051 3100 AY626231 (3051) TTCCACCCTGTTTACTTTCTTTCTTCAAATGTTACATAATAACATCCTGG TERT-507 (1) ---TCTTAATCCCCTTG-GA Consensus (3051) C TAAT C T G 3101 3150 AY626231 (3101) AGATTGGGCACAGGTACTATATACAGTGCTCTGGAATCCCACAGGGCTCC TERT-507 (17) AGATTGGGCACAGGTACTATATACAGTGCTCTGGAATCCCACAGGGCTCT Consensus (3101) AGATTGGGCACAGGTACTATATACAGTGCTCTGGAATCCCACAGGGCTC 3151 3200 AY626231 (3151) ATTTTGTCAACCTTACTTTGCAGCTTATGCTACGGAGACATGGAAAACAA TERT-507 (67) ATTTTGTCAACCTTACTTTGCAGCTTATGCTACGGAGACATGGAAAACAA Consensus (3151) ATTTTGTCAACCTTACTTTGCAGCTTATGCTACGGAGACATGGAAAACAA 3201 3250 AY626231 (3201) ATTACTCTGTGGGATCCAGAAGGATGGAGTCCTAATACGTCTTATTGATG TERT-507 (117) ATTACTCTGTGGGATCCAGAAGGATGGAGTCTTAATACGTCTTATTGATG Consensus (3201) ATTACTCTGTGGGATCCAGAAGGATGGAGTC TAATACGTCTTATTGATG 3251 3300 AY626231 (3251) ACTTTTTGCTGGTTACGCCACATTTAATGCAGGCAAGAACTTTTCTAAGG TERT-507 (167) ACTTTTTGCTGGTTACACCACATTTAATGCAGGCAAGAACTTTTCTAAGG Consensus (3251) ACTTTTTGCTGGTTAC CCACATTTAATGCAGGCAAGAACTTTTCTAAGG 3301 3350 AY626231 (3301) ACTATAGCAGCAGGTATTCCTGAGTATGGCTTTTTAATAAATGCCAAGAA TERT-507 (217) ACTATAGCAGCAGGTATTCCTGAGTATGGCTTTTTAATAAATGCCAAGAA Consensus (3301) ACTATAGCAGCAGGTATTCCTGAGTATGGCTTTTTAATAAATGCCAAGAA 3351 3400 AY626231 (3351) GACTGTGGTGAATTTTCCTGTTGATGATATCCCGGGATGTTCCAAGTTCA TERT-507 (267) GACGGTGGTGAATTTTCCTGTTGATGATATCCCGGGATGTTCCAAGTTCA Consensus (3351) GAC GTGGTGAATTTTCCTGTTGATGATATCCCGGGATGTTCCAAGTTCA 3401 3450 AY626231 (3401) AACATCTGCCAGATTGTCGTTTGATCTCATGGTGTGGTTTATTATTGGAT TERT-507 (317) AACATCTGCCAGATTGTCGCTTGATCTCATGGTGTGGTTTATTATTGGAT Consensus (3401) AACATCTGCCAGATTGTCG TTGATCTCATGGTGTGGTTTATTATTGGAT 3451 3500 AY626231 (3451) GTGCAGACACTTGAGGTTTATTGTGATTACTCCAGTTATGCCTTTACTTC TERT-507 (367) GTGCAGACACTTGAGGTTTATTGTGATTACTCCAGTTATGCCTTTACTTC Consensus (3451) GTGCAGACACTTGAGGTTTATTGTGATTACTCCAGTTATGCCTTTACTTC 3501 3550 AY626231 (3501) TATCAGATCAAGTCTTTCCTTCAATTCAAGTAGAATAGCTGGAAAAAACA TERT-507 (417) TATCAGATCAAGTCTTTCCTTCAATTCAAGTAGAATAGCTGGGAAAAACA Consensus (3501) TATCAGATCAAGTCTTTCCTTCAATTCAAGTAGAATAGCTGG AAAAACA 3551 3600 AY626231 (3551) TGAAATGCAAATTGACTGCAGTCCTCAAACTGAAATGCCATCCTTTACTT TERT-507 (467) TGAAATGCAAATTGA--- Consensus (3551) TGA

AATGCAAATTGA

圖 4. 白色來亨雞之 TERT基因產物進行 DNA序列分析與 Basic Local Alignment Search Tool (BLAST) 比對之結果

Fig. 4. DNA sequences and BLAST analysis of TERT gene

PCR-products amplified from gastrula stage embryos of White Leghorn chicken.

圖 5. 自白色來亨雞原腸期胚 (E) cDNA 進行 PCR 擴增的 TERT 基因前、中、後段基因。M為 1 kb 標記。

Fig. 5. Electrophoresis analysis of TERT PCR products amplified from gastrula stage embryos (E) in White leghorn chicken. M: 1 kp ladder marker.

Length 1700 bp (30570 bp to 4764 bp) Length 1209 bp

(2150 bp to 3358 bp) Length 1009 bp

(44 bp to 1052 bp)

Length 1203 bp (1001 bp to 2203 bp)

3.0 Kb 1.0 Kb

M 1 2

圖 6. 白色來亨雞 TERT 基因啟動子區 (1,307 bp) (1)與 3’ 端未轉 錄區 (609 bp) (2) 應用 PCR 選殖後獲得之產物。M為標記。

Fig. 6. Electrophoresis analysis of TERT promoter (1,307 bp) (1) and 3’

UTR (609 bp) (2) PCR products were amplified from gastrula stage embryos (E) in White leghorn chicken. M: ladder marker.

M E E Nc

3.0 kb 1.5 kb

4,799 bp

圖 7. 白色來亨雞 TERT全長基因之 PCR選殖產物 (4,799 bp)。M為 分子標記。Nc 為陰性對照。

Fig. 7. Full length of TERT PCR products (4,799 bp) amplified from gastrula stage embryos (E) in White Leghorn chicken. M: ladder marker. Na: negative control.

M XhoI/SacI

4.7 kb insert fragment 3.0 kb

1.5 kb

圖 8. XhoI/SacI切割篩選 pchTERT TOPO質體。4,799 bp片段顯示完 整的TERT基因。M為分子標記。

Fig. 8. The XhoI/SacI digested putative pchTERT TOPO. The 4,799 bp fragment indicated full length of TERT gene. M: ladder marker.

pchTERT TOPO 8418 bp

chTERT 3185-7979

TOPO vector 7980-3184

圖 9. 含有雞 TERT 基因的 pchTERT TOPO 序列分析載體圖譜 Fig. 9. The map of pchTERT TOPO.

圖 10. 進行 BLAST 比對後與白色來亨雞 TERT 之 DNA 具有相似性 的序列及相似位置圖

Fig. 10. The genes which share high similarity with the White Leghorn chicken TERT are indicated after BLAST alignment.

ref|NM_001031007.1| Gallus gallus telomerase reverse transcriptase

Query 1 ATGGAGCGCGGGGCTCAGCCGGGAGTCGGCGTGCGGCGGCTCCGCAATGTAGCGCGGGAG 60 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1 ATGGAGCGCGGGGCTCAGCCGGGAGTCGGCGTGCGGCGGCTCCGCAATGTAGCGCGGGAG 60

Query 61 GAGCCCTTCGCCGCGGTCCTGGGCGCGCTGCGGGGCTGCTACGCCGAGGCCACGCCGCTG 120 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 61 GAGCCCTTCGCCGCGGTCCTGGGCGCGCTGCGGGGCTGCTACGCCGAGGCCACGCCGCTG 120

Query 121 GAGGCCTTCGTCCGGCGGCTGCAGGAGGGTGGCACCGGGGAGGTCGAGGTGCTGCGAGGC 180 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 121 GAGGCCTTCGTCCGGCGGCTGCAGGAGGGTGGCACCGGGGAGGTCGAGGTGCTGCGAGGC 180

Query 181 GACGACGCTCAGTGCTACCGGACCTTCGTGTCGCAGTGCGTGGTGTGCGTCCCCCGCGGG 240 |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 181 GACGACGCTCAGTGCTACCGGACCTTCGTGTCGCAGTGCGTGGTGTGCGTCCCCCGCGGT 240

Query 241 GCTCGCGCCATCCCCCGGCCCATCTGCTTCCAGCAGTTATCCAGTCAGGGCGAAGTCATC 300 |||||||||||||||||||||||||||||||||||||||||||||||| |||||||||||

Sbjct 241 GCTCGCGCCATCCCCCGGCCCATCTGCTTCCAGCAGTTATCCAGTCAGAGCGAAGTCATC 300

Query 301 ACGAGAATCGTTCAGAGGCTGTGTGAAAAGAAAAAGAAGAACATCCTTGCGTATGGATAC 360 || |||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 301 ACAAGAATCGTTCAGAGGCTGTGTGAAAAGAAAAAGAAGAACATCCTTGCGTATGGATAC 360

Query 361 TCCTTGCTGGATGAGAACAGTTGTCACTTCAGAGTTTTGCCATCTTCGTGTATATACAGC 420 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 361 TCCTTGCTGGATGAGAACAGTTGTCACTTCAGAGTTTTGCCATCTTCGTGTATATACAGC 420

Query 421 TATCTGTCCAATACTGTAACAGAAACGATTCGCATCAGTGGCCTCTGGGAGATACTGCTG 480 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 421 TATCTGTCCAATACTGTAACAGAAACGATTCGCATCAGTGGCCTCTGGGAGATACTGCTG 480

Query 481 AGTAGGATAGGGGACGACGTGATGATGTACCTGCTGGAGCACTGTGCGCTCTTCATGCTG 540 ||||||||||||||||||||||||||||||||||||||||||||||| ||||||||||||

Sbjct 481 AGTAGGATAGGGGACGACGTGATGATGTACCTGCTGGAGCACTGTGCACTCTTCATGCTG 540

Query 541 GTTCCCCCAAGTAACTGTTACCAGGTCTGCGGGCAACCAATTTATGAACTTATTTCGCGT 600 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 541 GTTCCCCCAAGTAACTGTTACCAGGTCTGCGGGCAACCAATTTATGAACTTATTTCGCGT 600

Query 601 AACATAGGGCCATCCCCAGGGTTTGTTAGACGACGGTACTCAAGGTTTAAACATAATAGC 660 ||| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 601 AACGTAGGGCCATCCCCAGGGTTTGTTAGACGACGGTACTCAAGGTTTAAACATAATAGC 660

Query 661 TTGCTTGACTATGTGCGAAAAAGGCTTGTGTTTCACAGGCACTATCTTTCCAAGTCGCAA 720 |||||||||||||||||||||||||||||||||||||||||||||||||||||||| ||

Sbjct 661 TTGCTTGACTATGTGCGAAAAAGGCTTGTGTTTCACAGGCACTATCTTTCCAAGTCACAG 720

Query 721 TGGTGGAAGTGCAGGCCGAGACGTCGAGGTCGCGTCTCCAGCAGGAGAAAAAGAAGGAGC 780 |||||||||||||||||||||||||||||||| |||||||||||||||||||||||||||

Sbjct 721 TGGTGGAAGTGCAGGCCGAGACGTCGAGGTCGTGTCTCCAGCAGGAGAAAAAGAAGGAGC 780

Query 781 CATAGGATACAAAGCCTAAGGTCTGGTTATCAGCCTTCTGCAAAAGTGAACTTTCAAGCA 840 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 781 CATAGGATACAAAGCCTAAGGTCTGGTTATCAGCCTTCTGCAAAAGTGAACTTTCAAGCA 840

Query 841 GGTAGGCAGATCAGCACTGTTACTGCACGTCTGGAAAAACAGAGCTGCTCCAGTTTATGT 900 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 841 GGTAGGCAGATCAGCACTGTTACTGCACGTCTGGAAAAACAGAGCTGCTCCAGTTTATGT 900

Query 901 TTGCCAGCTAGAGCACCATCTTTAAAAAGGAAGCGTGATGGAGAACAGGTTGAAATCACA 960 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 901 TTGCCAGCTAGAGCACCATCTTTAAAAAGGAAGCGTGATGGAGAACAGGTTGAAATCACA 960

Query 961 GCTAAGAGAGTGAAAGTAATGGAGAAAGAGATAGAGGAACAGGCTTGTAGTATCGTTCCT 1020 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 961 GCTAAGAGAGTGAAAGTAATGGAGAAAGAGATAGAGGAACAGGCTTGTAGTATCGTTCCT 1020

Query 1021 GATGTAAACCAAAGTAGCTCCCAGAGGCATGGAACCTCCTGGCATGTAGCACCACGTGCT 1080 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1021 GATGTAAACCAAAGTAGCTCCCAGAGGCATGGAACCTCCTGGCATGTAGCACCACGTGCT 1080

Query 1081 GTAGGTCTTATTAAAGAACATTACATTTCTGAAAGAAGTAACAGTGAGATGTCTGGTCCT 1140 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1081 GTAGGTCTTATTAAAGAACATTACATTTCTGAAAGAAGTAACAGTGAGATGTCTGGTCCT 1140

Query 1141 TCTGTAGTTCGCAGATCTCACCCTGGGAAGAGGCCTGTGGCAGACAAAAGCTCTTTTCCA 1200 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1141 TCTGTAGTTCGCAGATCTCACCCTGGGAAGAGGCCTGTGGCAGACAAAAGCTCTTTTCCA 1200

Query 1201 CAAGGAGTTCAGGGTAACAAACGCATAAAGACCGGTGCAGAAAAACGAGCAGAATCCAAT 1260 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1201 CAAGGAGTTCAGGGTAACAAACGCATAAAGACCGGTGCAGAAAAACGAGCAGAATCCAAT 1260

Query 1261 AGAAGGGGCATAGAGATGTATATAAACCCAATCCATAAACCCAATAGAAGGGGCATAGAG 1320 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1261 AGAAGGGGCATAGAGATGTATATAAACCCAATCCATAAACCCAATAGAAGGGGCATAGAG 1320

Query 1321 AGGCGTATAAATCCAACCCACAAACCTGAGTTGAATTCTGTACAAACTGAACCAATGGAA 1380 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1321 AGGCGTATAAATCCAACCCACAAACCTGAGTTGAATTCTGTACAAACTGAACCAATGGAA 1380

Query 1381 GGTGCTTCTTCAGGGGACAGAAAGCAGGAAAATCCCCCAGCTCATTTGGCAAAGCAGTTA 1440 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1381 GGTGCTTCTTCAGGGGACAGAAAGCAGGAAAATCCCCCAGCTCATTTGGCAAAGCAGTTA 1440

Query 1441 CCAAATACATTGTCGCGCTCTACAGTGTACTTTGAGAAGAAATTTCTTCTGTATTCCCGC 1500 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1441 CCAAATACATTGTCGCGCTCTACAGTGTACTTTGAGAAGAAATTTCTTCTGTATTCCCGC 1500

Query 1501 AGTTACCAAGAATATTTTCCTAAATCGTTCATACTGAGCCGCCTGCAGGGTTGTCAGGCA 1560 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1501 AGTTACCAAGAATATTTTCCTAAATCGTTCATACTGAGCCGCCTGCAGGGTTGTCAGGCA 1560

Query 1561 GGTGGAAGGCGGCTTATAGAAACTATATTCTTAAGCCAAAACCCATTAAAGGAACAGCAG 1620 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1561 GGTGGAAGGCGGCTTATAGAAACTATATTCTTAAGCCAAAACCCATTAAAGGAACAGCAG 1620

Query 1621 AACCAAAGCCTACCACAGCAAAAGTGGCGAAAGAAGAGGTTGCCCAAACGCTACTGGCAA 1680 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1621 AACCAAAGCCTACCACAGCAAAAGTGGCGAAAGAAGAGGTTGCCCAAACGCTACTGGCAA 1680

Query 1681 ATGAGAGAGATATTTCAGAAGCTGGTAAAGAACCATGAGAAGTGCCCTTATTTAGTTTTC 1740 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1681 ATGAGAGAGATATTTCAGAAGCTGGTAAAGAACCATGAGAAGTGCCCTTATTTAGTTTTC 1740

Query 1741 TTGAGGAAAAATTGCCCTGTTTTGCTTTCTGAAGCATGTTTGAAAAAGACGGAGCTGACC 1800 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1741 TTGAGGAAAAATTGCCCTGTTTTGCTTTCTGAAGCATGTTTGAAAAAGACGGAGCTGACC 1800

Query 1801 TTGCAGGCGGCTCTGCCTGGGGAAGCAAAGGTTCACAAGCACACAGAACATGGGAAAGAG 1860 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1801 TTGCAGGCGGCTCTGCCTGGGGAAGCAAAGGTTCACAAGCACACAGAACATGGGAAAGAG 1860

Query 1861 TCCACTGAGGGTACTGCACCGAACAGCTTCCTCGCTCCTCCCTCAGTGCTAGCGTGTGGG 1920 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1861 TCCACTGAGGGTACTGCACCGAACAGCTTCCTCGCTCCTCCCTCAGTGCTAGCGTGTGGG 1920

Query 1921 CAGCCAGAGAGAGGGGAACAGCACCCTGCAGAGGGGAGTGATCCGCTCCTCAGGGAGCTG 1980 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1921 CAGCCAGAGAGAGGGGAACAGCACCCTGCAGAGGGGAGTGATCCGCTCCTCAGGGAGCTG 1980

Query 1981 CTCAGGCAGCACAGCAGCCACTGGCAGGTGTATGGCTTTGTGAGGGAGTGCCTGGAGCGG 2040 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 1981 CTCAGGCAGCACAGCAGCCACTGGCAGGTGTATGGCTTTGTGAGGGAGTGCCTGGAGCGG 2040

Query 2041 GTGATCCCTGCTGAGCTGTGGGGTTCAAGCCATAACAAATGCCGGTTCTTTAAAAACGTG 2100 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2041 GTGATCCCTGCTGAGCTGTGGGGTTCAAGCCATAACAAATGCCGGTTCTTTAAAAACGTG 2100

Query 2101 AAAGCATTCATTTCCATGGGGAAGTATGCTAAGCTTTCATTGCAGCAGCTGATGTGGAAG 2160 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2101 AAAGCATTCATTTCCATGGGGAAGTATGCTAAGCTTTCATTGCAGCAGCTGATGTGGAAG 2160

Query 2161 ATGAGAGTGAATGACTGCGTATGGCTTCGTCTGGCCAAAGGTAATCACTCTGTTCCTGCC 2220 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2161 ATGAGAGTGAATGACTGCGTATGGCTTCGTCTGGCCAAAGGTAATCACTCTGTTCCTGCC 2220

Query 2221 TATGAACATTGTTACCGTGAAGAAATTCTGGCAAAATTCCTATACTGGCTGATGGATTCC 2280 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2221 TATGAACATTGTTACCGTGAAGAAATTCTGGCAAAATTCCTATACTGGCTGATGGATTCC 2280

Query 2281 TATGTTATCGAGTTGCTCAAATCATTTTTCTATATCACCGAGACCATGTTCCAGAAAAAC 2340 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2281 TATGTTATCGAGTTGCTCAAATCATTTTTCTATATCACCGAGACCATGTTCCAGAAAAAC 2340

Query 2341 ATGCTTTTCTACTACCGAAAGTTTATCTGGGGCAAGTTACAGAACATTGGAATTAGAGAC 2400 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2341 ATGCTTTTCTACTACCGAAAGTTTATCTGGGGCAAGTTACAGAACATTGGAATTAGAGAC 2400

Query 2401 CATTTTGCCAAAGTACATCTACGTGCCTTGTCTTCAGAGGAGATGGAAGTGATCCGTCaa 2460 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2401 CATTTTGCCAAAGTACATCTACGTGCCTTGTCTTCAGAGGAGATGGAAGTGATCCGTCAA 2460

Query 2461 aaaaaGTATTTTCCTATTGCATCAAGGCTCCGGTTCATTCCTAAAATGAATGGTTTAAGA 2520 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2461 AAAAAGTATTTTCCTATTGCATCAAGGCTCCGGTTCATTCCTAAAATGAATGGTTTAAGA 2520

Query 2521 CCCGTAGTAAGACTAAGCCGTGTTGTTGAAGGACAGAAACTCAGCAAGGAAAGCAGAGAA 2580 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2521 CCCGTAGTAAGACTAAGCCGTGTTGTTGAAGGACAGAAACTCAGCAAGGAAAGCAGAGAA 2580

Query 2581 AAGAAGATACAGCGCTATAACACTCAGCTAAAAAATCTATTTAGTGTTTTAAACTATGAA 2640 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2581 AAGAAGATACAGCGCTATAACACTCAGCTAAAAAATCTATTTAGTGTTTTAAACTATGAA 2640

Query 2641 CGAACTGTAAACACCAGTATCATTGGCTCTTCAGTATTCGGGAGAGATGATATCTACAGG 2700 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2641 CGAACTGTAAACACCAGTATCATTGGCTCTTCAGTATTCGGGAGAGATGATATCTACAGG 2700

Query 2701 AAGTGGAAGGAGTTTGTTACAAAGGTTTTTGAATCAGGTGGTGAAATGCCTCATTTCTAC 2760 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2701 AAGTGGAAGGAGTTTGTTACAAAGGTTTTTGAATCAGGTGGTGAAATGCCTCATTTCTAC 2760

Query 2761 TTTGTAAAGGGTGATGTATCCAGAGCTTTTGATACCATTCCTCACAAGAAACTTGTGGAA 2820 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2761 TTTGTAAAGGGTGATGTATCCAGAGCTTTTGATACCATTCCTCACAAGAAACTTGTGGAA 2820

Query 2821 GTGATATCACAGGTCTTGAAACCTGAGAGCCAAACTGTCTATGGAATAAGGTGGTATGCA 2880 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2821 GTGATATCACAGGTCTTGAAACCTGAGAGCCAAACTGTCTATGGAATAAGGTGGTATGCA 2880

Query 2881 GTGATTATGATTACCCCAACTGGAAAAGCCAGGAAACTCTATAAGAGACATGTTTCTACT 2940 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2881 GTGATTATGATTACCCCAACTGGAAAAGCCAGGAAACTCTATAAGAGACATGTTTCTACT 2940

Query 2941 TTCGAGGATTTTATTCCAGACATGAAGCAGTTTGTGTCCAAGCTTCAAGAGAGAACTTCA 3000 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 2941 TTCGAGGATTTTATTCCAGACATGAAGCAGTTTGTGTCCAAGCTTCAAGAGAGAACTTCA 3000

Query 3001 TTACGAAATGCAATAGTAGTTGAACAGTGCTTAACTTTTAATGAGAACAGTTCCACCCTG 3060

||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3001 TTACGAAATGCAATAGTAGTTGAACAGTGCTTAACTTTTAATGAGAACAGTTCCACCCTG 3060

Query 3061 TTTACTTTCTTTCTTCAAATGTTACATAATAACATCCTGGAGATTGGGCACAGGTACTAT 3120 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3061 TTTACTTTCTTTCTTCAAATGTTACATAATAACATCCTGGAGATTGGGCACAGGTACTAT 3120

Query 3121 ATACAGTGCTCTGGAATCCCACAGGGCTCCATTTTGTCAACCTTACTTTGCAGCTTATGC 3180 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3121 ATACAGTGCTCTGGAATCCCACAGGGCTCCATTTTGTCAACCTTACTTTGCAGCTTATGC 3180

Query 3181 TACGGAGACATGGAAAACAAATTACTCTGTGGGATCCAGAAGGATGGAGTCCTAATACGT 3240 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3181 TACGGAGACATGGAAAACAAATTACTCTGTGGGATCCAGAAGGATGGAGTCCTAATACGT 3240

Query 3241 CTTATTGATGACTTTTTGCTGGTTACGCCACATTTAATGCAGGCAAGAACTTTTCTAAGG 3300 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3241 CTTATTGATGACTTTTTGCTGGTTACGCCACATTTAATGCAGGCAAGAACTTTTCTAAGG 3300

Query 3301 ACTATAGCAGCAGGTATTCCTGAGTATGGCTTTTTAATAAATGCCAAGAAGACTGTGGTG 3360 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3301 ACTATAGCAGCAGGTATTCCTGAGTATGGCTTTTTAATAAATGCCAAGAAGACTGTGGTG 3360

Query 3361 AATTTTCCTGTTGATGATATCCCGGGATGTTCCAAGTTCAAACATCTGCCAGATTGTCGT 3420 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3361 AATTTTCCTGTTGATGATATCCCGGGATGTTCCAAGTTCAAACATCTGCCAGATTGTCGT 3420

Query 3421 TTGATCTCATGGTGTGGTTTATTATTGGATGTGCAGACACTTGAGGTTTATTGTGATTAC 3480 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3421 TTGATCTCATGGTGTGGTTTATTATTGGATGTGCAGACACTTGAGGTTTATTGTGATTAC 3480

Query 3481 TCCAGTTATGCCTTTACTTCTATCAGATCAAGTCTTTCCTTCAATTCAAGTAGAATAGCT 3540 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3481 TCCAGTTATGCCTTTACTTCTATCAGATCAAGTCTTTCCTTCAATTCAAGTAGAATAGCT 3540

Query 3541 GGGAAAAACATGAAATGCAAATTGACTGCAGTCCTCAAACTGAAATGCCATCCTTTACTT 3600 || |||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3541 GGAAAAAACATGAAATGCAAATTGACTGCAGTCCTCAAACTGAAATGCCATCCTTTACTT 3600

Query 3601 CTTGACTTAAAGATCAACAGCCTTCAGACAGTTCTAATTAACATCTACAAGATATTTTTA 3660 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3601 CTTGACTTAAAGATCAACAGCCTTCAGACAGTTCTAATTAACATCTACAAGATATTTTTA 3660

Query 3661 CTTCAGGCTTACAGGTTCCATGCCTGTGTTCTTCAGCTTCCATTCAACCAGAAAGTTAGG 3720 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3661 CTTCAGGCTTACAGGTTCCATGCCTGTGTTCTTCAGCTTCCATTCAACCAGAAAGTTAGG 3720

Query 3721 AATAATCCTGATTTCTTCCTAAGGATCATCTCTGATACTGCTTCATGCTGCTATTTTATC 3780 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3721 AATAATCCTGATTTCTTCCTAAGGATCATCTCTGATACTGCTTCATGCTGCTATTTTATC 3780

Query 3781 CTGAAAGCTAAAAATCCAGGAGTTTCTTTAGGTAGCAAGGATGCATCTGGCATGTTCCCT 3840 |||||||||||||||||||||||||||||||||||||| |||||||||||||||||||||

Sbjct 3781 CTGAAAGCTAAAAATCCAGGAGTTTCTTTAGGTAGCAAAGATGCATCTGGCATGTTCCCT 3840

Query 3841 TTTGAGGCAGCAGAATGGCTGTGCTACCATGCCTTCATTGTCAAACTGTCCAACCACAAA 3900 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3841 TTTGAGGCAGCAGAATGGCTGTGCTACCATGCCTTCATTGTCAAACTGTCCAACCACAAA 3900

Query 3901 GTTATTTACAAATGCTTACTTAAGCCCCTTAAAGTCTATAAGATGCGTCTGTTTGGGAAG 3960 |||||||||||||||||||||||||||||||||||||||||||||| |||||||||||||

Sbjct 3901 GTTATTTACAAATGCTTACTTAAGCCCCTTAAAGTCTATAAGATGCATCTGTTTGGGAAG 3960

Query 3961 ATCCCAAGGGATACTATGGAACTGCTGAAGACGGTGACGGAACCATCGCTTTGTCAAGAT 4020 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 3961 ATCCCAAGGGATACTATGGAACTGCTGAAGACGGTGACGGAACCATCGCTTTGTCAAGAT 4020

Query 4021 TTCAAAACTATACTGGACTAAGGGATGAAGGTCTATTTCATAATCCTTCCAGGCTTAAAA 4080 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4021 TTCAAAACTATACTGGACTAAGGGATGAAGGTCTATTTCATAATCCTTCCAGGCTTAAAA 4080

Query 4081 ATAAATCTTAAATTTTTAGATTAATTTTCTCTGTAGATAATTTTGGTAGACTGAACTGAA 4140 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4081 ATAAATCTTAAATTTTTAGATTAATTTTCTCTGTAGATAATTTTGGTAGACTGAACTGAA 4140

Query 4141 AGCATCTCTACTTGAAAGTTTTCtttttttttttATGGGTACCACACAATGACCTTTAAA 4200 ||||||||||||||||||||||||||||||||||||||||||||| ||||||||||||||

Sbjct 4141 AGCATCTCTACTTGAAAGTTTTCTTTTTTTTTTTATGGGTACCACCCAATGACCTTTAAA 4200

Query 4201 AGAGTAGGCTGTTTTGTTAACATCTTCATGTTTGTGTTTTTCTGTAAGTTCGAAGGCTGT 4260 ||||||| ||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4201 AGAGTAGACTGTTTTGTTAACATCTTCATGTTTGTGTTTTTCTGTAAGTTCGAAGGCTGT 4260

Query 4261 TTGAAGTACAACCTGTGAACTTGACTTGATTTCCTTCAGTAATTCCTTTCTTCTTGGTGA 4320 |||||||||||||||||||||||||| |||||||||||||||||||||||||||||||||

Sbjct 4261 TTGAAGTACAACCTGTGAACTTGACTCGATTTCCTTCAGTAATTCCTTTCTTCTTGGTGA 4320

Query 4321 AGACATATTTAACCTATCTGTATAAACAGATCAATATTTTTGCCAGTATTGTTTCTGAAG 4380 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4321 AGACATATTTAACCTATCTGTATAAACAGATCAATATTTTTGCCAGTATTGTTTCTGAAG 4380

Query 4381 AACTGTTCTTTGATTTTTCTATTGCAGACAATTTTACAATAAATTGTCTACATAGTTGCA 4440 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4381 AACTGTTCTTTGATTTTTCTATTGCAGACAATTTTACAATAAATTGTCTACATAGTTGCA 4440

Query 4441 ATAATGCATACTTGCCTCCAATACCTTTATTTTGTAGTAAGCTTCAACCCAATCCCACCC 4500 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4441 ATAATGCATACTTGCCTCCAATACCTTTATTTTGTAGTAAGCTTCAACCCAATCCCACCC 4500

Query 4501 AACTTGGCAGTGTCTTGAAATGCACTTATTGGTACAGGGAAAGTGCATTTCATGGCTGCG 4560 ||||||||| ||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4501 AACTTGGCATTGTCTTGAAATGCACTTATTGGTACAGGGAAAGTGCATTTCATGGCTGCG 4560

Query 4561 CCTAGCAAGATATGTGGGGTAAAGATTCTTTCACTTTGGAGAACACAGTCAAAATTTGTA 4620 ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||

Sbjct 4561 CCTAGCAAGATATGTGGGGTAAAGATTCTTTCACTTTGGAGAACACAGTCAAAATTTGTA 4620

在文檔中 嘉南藥理科技大學 生物科技系 碩士論文 (頁 56-106)