中国农业科学 ›› 2019, Vol. 52 ›› Issue (21): 3713-3732.doi: 10.3864/j.issn.0578-1752.2019.21.001
收稿日期:
2019-04-29
接受日期:
2019-06-18
出版日期:
2019-11-01
发布日期:
2019-11-12
通讯作者:
贺道华
作者简介:
张奇艳,E-mail: 基金资助:
ZHANG QiYan1,LEI ZhongPing2,SONG Yin1,HAI JiangBo1,HE DaoHua1()
Received:
2019-04-29
Accepted:
2019-06-18
Online:
2019-11-01
Published:
2019-11-12
Contact:
DaoHua HE
摘要:
目的 扩展蛋白(Expansin)是细胞壁的重要组成部分,在植物的生长发育及逆境胁迫应答等方面均发挥着重要作用。基于全基因组水平系统鉴定陆地棉Expansin基因家族,并通过生物信息学及表达模式分析,为揭示扩展蛋白基因在棉花生长发育中的功能及后续利用奠定基础。方法 利用BLAST和HMMER在陆地棉基因组中搜索并鉴定扩展蛋白基因家族成员;利用ClustalW、MEGA、MCScanX、Prot Param、MEME、SignalP、Euk-mPLoc、FancyGene和DnaSP等软件对其基因序列和蛋白序列进行生物信息学分析。通过RNA-seq数据分析扩展蛋白基因的表达模式和部分同源基因间表达差异,利用qRT-PCR验证部分扩展蛋白基因的表达谱。结果 陆地棉基因组中含有46个EXPA基因、8个EXPB基因、6个EXLA基因和12个EXLB基因,合计72个Expansin基因;四倍体陆地棉中扩展蛋白成员的数量几乎是二倍体棉种(亚洲棉与雷蒙德氏棉)的2倍。除GhA02和GhD06 2个染色体外,其余各染色体上均分布有数目不等的扩展蛋白基因(2—4个),具有部分同源关系的染色体GhA08和GhD08分别有5个和8个扩展蛋白基因。系统发育树显示,各亚家族成员聚集成群,并且大部分的末端分支均由来源于3个物种的4个(亚)基因组的4个基因组成,如EXPA亚家族的Cotton_A_28454/Gh_A03G0885/Gh_D02G1269/Gorai. 005G142200等等,4个基因之间具有同线性关系。亚细胞定位发现陆地棉所有的扩展蛋白均位于细胞外。基因结构分析显示,扩展蛋白基因由3—5个外显子组成,外显子-内含子结构在进化上高度保守且与氨基酸序列的多样性一致,且在外显子上存在密码子偏好性。RNA-seq数据显示,不同基因在不同时空条件下存在特异性表达,如GhEXPA19A和GhEXPA19D相比其他基因在纤维10 DPA和20 DPA中的表达量很高;在不同的组织(如子叶、新叶、老叶、苞叶)中,GhEXPA24D具有较高的表达量。部分同源基因之间具有不同的表达模式,显示它们之间功能的异化与互补。qRT-PCR结果与RNA-seq数据基本吻合,如GhEXLA3A和GhEXLA3D在纤维发育的伸长阶段高量表达。GhEXPA19D和GhEXLA2D在3DPA的胚珠中表达活跃。结论 陆地棉基因组中含72个扩展蛋白基因,其在DNA水平和氨基酸水平具有一致的结构多样性和进化保守性,在转录水平具有各异的表达模式,显示出家族内成员间功能上的异化与互补。
张奇艳, 雷忠萍, 宋银, 海江波, 贺道华. 陆地棉扩展蛋白基因的鉴定与特征分析[J]. 中国农业科学, 2019, 52(21): 3713-3732.
ZHANG QiYan, LEI ZhongPing, SONG Yin, HAI JiangBo, HE DaoHua. Identification and Characterization of the Expansin Gene Family in Upland Cotton (Gossypium hirsutum)[J]. Scientia Agricultura Sinica, 2019, 52(21): 3713-3732.
表1
实时荧光定量PCR引物"
基因名称Gene name | 正向引物Forward primers (5′-3′) | 反向引物Reverse primers (5′-3′) |
---|---|---|
GhEXLA1A | TCACTGACATTGCAAAAGAAGG | GAACACCATTGATTGTGTAGCA |
GhEXLA3A | GTGATTACAGCCATTGGTGATG | CCAAAACAAAAGCTCTCAAAGACTC |
GhEXLA2D | TAACATGCAAAGAAAGTGAGGC | AATCCATGGTTTTCCGGAAATG |
GhEXLA3D | CTTTCGGGTCACTCAAAAATGT | AATGCTTCTCAACATCCCAAAC |
GhEXPA06A | CTTTCGGGTCACTCAAAAATGT | AATGCTTCTCAACATCCCAAAC |
GhEXPA16A | GAATCGTCCCCATTTCATTCAG | CACCACCAACATTTGTAGTGAG |
GhEXPA25A | CACCATCATTGTAACTGCTACG | ATTTTGAACTTAACACCACCGG |
GhEXPA06D | GCACTGCTTTGTTTAACAATGG | ATTAGGGAGAGCATTGTTTGGA |
GhEXPA17D | TTGTTGGGTTTTCTAGCAATGG | CTGCAGTATTTGTTCCATAGCC |
GhEXPA19D | GGAAACCTGTACAGTCAAGGTA | GTTATGGTTCGACTAATGCACC |
GhEXPA20D | CCTAACTTAGCATTGTCCAACG | TTTCTTCATGCAGGGTACTCTT |
GhEXPA23D | CGGCCACTCATACTTTAACTTG | CTTCGTATGTTTGTCCGAACTG |
GhEXPA24D | GAACGAAGAAAACACCCATCTT | CTCTTCAGCCAAAGCATCTTAC |
UBQ7 | GAAGGCATTCCACCTGACCAAC | CTTGACCTTCTTCTTCTTGTGCTTG |
表2
陆地棉扩展蛋白基因家族成员的鉴定"
基因 Gene | 序列号 Phytozome ID | 染色体位置 Location | 基因长度Length of gene (bp) | 外显子个数Number of exon | 氨基酸个数 Number of amino acids | 信号 肽长度Signal peptide (aa) | 分子量Molecular weight (kD) | 理论等电点Theoretical pI | 蛋白不稳定指数Instability index | 脂溶指数Aliphatic index | 总平均 疏水性GRAVY |
---|---|---|---|---|---|---|---|---|---|---|---|
GhEXPA03A | Gh_A01G0477 | GhA01: 7603263..7604881(+) | 1619 | 3 | 256 | 27 | 28.40 | 9.90 | 43.09 | 67.85 | -0.118 |
GhEXPA06A | Gh_A03G0885 | GhA03: 56794871..56796095(-) | 1225 | 3 | 248 | 20 | 26.59 | 9.30 | 32.13 | 62.94 | -0.140 |
GhEXPA16A | Gh_A05G2385 | GhA05: 29485491..29486507(-) | 1017 | 3 | 253 | 25 | 26.94 | 7.53 | 30.72 | 59.8 | -0.166 |
GhEXPA22-1A | Gh_A05G3210 | GhA05: 83889906..83890791(+) | 886 | 2 | 262 | 22 | 29.18 | 9.25 | 20.09 | 68.47 | -0.167 |
GhEXPA22-2A | Gh_A05G3211 | GhA05: 83905450..83906383(+) | 934 | 2 | 278 | 23 | 30.96 | 9.24 | 18.98 | 74.64 | 0.008 |
GhEXPA21A | Gh_A05G3493 | GhA05: 90545166..90546162(+) | 997 | 3 | 253 | 25 | 27.07 | 6.78 | 31.44 | 65.57 | -0.066 |
GhEXPA14A | Gh_A06G0018 | GhA06: 82222..83190(+) | 969 | 3 | 255 | 23 | 27.38 | 9.15 | 26.88 | 67.76 | -0.015 |
GhEXPA01A | Gh_A07G1208 | GhA07: 26997118..26998121(-) | 1004 | 3 | 258 | NO | 27.92 | 9.70 | 38.52 | 65.81 | -0.140 |
GhEXPA02A | Gh_A07G1881 | GhA07: 74202971..74204340(-) | 1370 | 3 | 253 | 21 | 27.52 | 9.40 | 38.41 | 77.79 | -0.025 |
GhEXPA05A | Gh_A08G1595 | GhA08: 94460350..94461323(+) | 974 | 3 | 267 | NO | 29.21 | 8.38 | 30.56 | 65.43 | 0.045 |
GhEXPA07A | Gh_A09G0135 | GhA09: 3242057..3243766(+) | 1710 | 3 | 256 | 21 | 28.16 | 9.22 | 41.30 | 68.52 | -0.160 |
GhEXPA08A | Gh_A09G1727 | GhA09: 70679591..70680590(-) | 1000 | 3 | 265 | 28 | 28.73 | 8.59 | 32.02 | 60.79 | -0.005 |
GhEXPA20A | Gh_A10G1606 | GhA10: 86603436..86604481(-) | 1046 | 3 | 253 | 25 | 27.00 | 7.55 | 33.15 | 62.89 | -0.118 |
GhEXPA19A | Gh_A10G2323 | scaffold2672_A10: 149124..150059(-) | 936 | 3 | 258 | 30 | 27.94 | 8.36 | 29.01 | 62.83 | -0.090 |
GhEXPA09A | Gh_A11G0187 | GhA11: 1736060..1737333(+) | 1274 | 4 | 265 | NO | 29.26 | 9.42 | 29.16 | 66.26 | -0.040 |
GhEXPA10A | Gh_A11G0964 | GhA11: 10482352..10483322(-) | 971 | 3 | 258 | NO | 28.42 | 9.43 | 33.85 | 66.98 | -0.229 |
GhEXPA11A | Gh_A11G2155 | GhA11: 71713430..71714457(-) | 1028 | 3 | 247 | 20 | 26.39 | 9.32 | 38.00 | 69.55 | -0.027 |
GhEXPA13A | Gh_A12G1619 | GhA12: 77299134..77300408(-) | 1275 | 3 | 248 | 20 | 26.62 | 9.19 | 31.75 | 69.6 | -0.030 |
GhEXPA23A | Gh_A13G0050 | GhA13: 540164..541646(+) | 1483 | 3 | 254 | 26 | 27.28 | 8.41 | 33.34 | 63.39 | -0.134 |
GhEXPA24A | Gh_A13G0672 | GhA13: 20543406..20544365(-) | 960 | 3 | 248 | 17 | 26.26 | 9.28 | 32.55 | 67.3 | -0.111 |
GhEXPA25A | Gh_A13G0719 | GhA13: 23714615..23715449(+) | 835 | 2 | 254 | 27 | 28.23 | 8.33 | 27.99 | 75.24 | -0.061 |
GhEXPA26A | Gh_A13G2059 | GhA13: 79891203..79892227(+) | 1025 | 3 | 249 | 23 | 26.93 | 8.36 | 29.07 | 68.59 | -0.064 |
GhEXPA03D | Gh_D01G0492 | GhD01: 5972132..5973758(+) | 1627 | 3 | 252 | NO | 27.98 | 9.90 | 39.24 | 70.48 | -0.078 |
GhEXPA06D | Gh_D02G1269 | GhD02: 41719506..41720746(-) | 1241 | 3 | 267 | NO | 28.95 | 9.78 | 35.24 | 68.31 | -0.104 |
基因 Gene | 序列号 Phytozome ID | 染色体位置 Location | 基因长度Length of gene (bp) | 外显子个数Number of exon | 氨基酸个数 Number of amino acids | 信号 肽长度Signal peptide (aa) | 分子量Molecular weight (kD) | 理论等电点Theoretical pI | 蛋白不稳定指数Instability index | 脂溶指数Aliphatic index | 总平均 疏水性GRAVY |
GhEXPA22D | Gh_D04G0396 | GhD04: 6280383..6281268(-) | 886 | 2 | 262 | 23 | 29.21 | 9.18 | 20.03 | 68.47 | -0.143 |
GhEXPA21D | Gh_D04G1924 | scaffold3943_D04: 15458..16477(-) | 1020 | 3 | 253 | 25 | 27.08 | 8.02 | 32.54 | 65.57 | -0.065 |
GhEXPA15D | Gh_D05G2433 | GhD05: 24370173..24371133(+) | 961 | 3 | 255 | 23 | 27.45 | 8.88 | 29.40 | 67.02 | -0.077 |
GhEXPA16D | Gh_D05G2650 | GhD05: 27568523..27569536(-) | 1014 | 3 | 252 | 24 | 26.92 | 7.53 | 30.80 | 60.79 | -0.130 |
GhEXPA17D | Gh_D05G2934 | GhD05: 34837918..34838746(-) | 829 | 2 | 253 | 29 | 27.21 | 9.04 | 31.30 | 64.78 | -0.122 |
GhEXPA01D | Gh_D07G1309 | GhD07: 20886446..20887308(-) | 863 | 3 | 207 | NO | 22.54 | 9.90 | 35.72 | 67.83 | -0.247 |
GhEXPA02D | Gh_D07G2095 | GhD07: 51074809..51076181(-) | 1373 | 3 | 253 | 21 | 27.49 | 9.30 | 38.37 | 76.25 | -0.044 |
GhEXPA04D | Gh_D08G1362 | GhD08: 44540800..44541775(-) | 976 | 254 | 23 | 28.01 | 9.31 | 35.94 | 67.68 | -0.187 | |
GhEXPA05D | Gh_D08G1905 | GhD08: 56979061..56980034(+) | 974 | 3 | 267 | NO | 29.32 | 8.08 | 29.33 | 66.14 | 0.037 |
GhEXPA07D | Gh_D09G0128 | GhD09: 3266295..3268335(+) | 2041 | 4 | 303 | NO | 33.80 | 9.57 | 43.94 | 72.38 | -0.164 |
GhEXPA08D | Gh_D09G1840 | GhD09: 45961349..45962346(-) | 998 | 3 | 265 | 28 | 28.74 | 8.38 | 34.27 | 59.32 | -0.006 |
GhEXPA19D | Gh_D10G1145 | GhD10: 18608318..18609246(+) | 929 | 3 | 258 | 30 | 27.95 | 8.05 | 29.69 | 62.44 | -0.106 |
GhEXPA20D | Gh_D10G1861 | GhD10: 52037144..52038189(-) | 1046 | 3 | 253 | 25 | 27.07 | 7.55 | 33.33 | 65.97 | -0.085 |
GhEXPA09D | Gh_D11G0198 | GhD11: 1728429..1729700(+) | 1272 | 4 | 265 | NO | 29.17 | 9.41 | 28.62 | 64.83 | -0.051 |
GhEXPA10D | Gh_D11G1115 | GhD11: 10243558..10244528(-) | 971 | 3 | 258 | NO | 28.42 | 9.43 | 33.85 | 66.98 | -0.229 |
GhEXPA11D | Gh_D11G2459 | GhD11: 49988580..49989605(-) | 1026 | 3 | 247 | 20 | 26.35 | 9.30 | 36.98 | 69.15 | -0.039 |
GhEXPA13D | Gh_D12G1759 | GhD12: 49966434..49967705(-) | 1272 | 3 | 248 | 20 | 26.77 | 9.19 | 34.40 | 69.19 | -0.016 |
GhEXPA23D | Gh_D13G0060 | GhD13: 622156..623599(+) | 1444 | 3 | 254 | 26 | 27.36 | 8.61 | 29.33 | 58.78 | -0.198 |
GhEXPA24D | Gh_D13G0786 | GhD13: 13599569..13600499(-) | 931 | 3 | 248 | 17 | 26.33 | 9.30 | 31.82 | 65.69 | -0.111 |
GhEXPA25D | Gh_D13G0843 | GhD13: 15386343..15387204(+) | 862 | 2 | 254 | 27 | 28.19 | 8.53 | 28.55 | 73.7 | -0.042 |
GhEXPA26D | Gh_D13G2460 | GhD13: 60483670..60484694(+) | 1025 | 3 | 249 | 23 | 26.86 | 8.36 | 29.68 | 68.59 | -0.067 |
GhEXPA17A | Gh_Sca005793G01 | scaffold5793: 5707..6535(+) | 829 | 2 | 253 | 29 | 27.17 | 8.88 | 29.73 | 68.62 | -0.076 |
GhEXPB2A | Gh_A03G2139 | scaffold729_A03: 85394..86864(+) | 1471 | 4 | 267 | 26 | 29.31 | 8.63 | 35.29 | 75.96 | -0.247 |
GhEXPB1A | Gh_A08G1231 | GhA08: 82800735..82801892(-) | 1158 | 4 | 273 | NO | 28.60 | 5.19 | 37.99 | 70.07 | 0.000 |
GhEXPB3A | Gh_A12G0861 | GhA12: 57318819..57321212(-) | 2394 | 5 | 256 | NO | 26.80 | 4.88 | 35.29 | 74.69 | -0.083 |
基因 Gene | 序列号 Phytozome ID | 染色体位置 Location | 基因长度Length of gene (bp) | 外显子个数Number of exon | 氨基酸个数 Number of amino acids | 信号 肽长度Signal peptide (aa) | 分子量Molecular weight (kD) | 理论等电点Theoretical pI | 蛋白不稳定指数Instability index | 脂溶指数Aliphatic index | 总平均 疏水性GRAVY |
GhEXPB4A | Gh_A12G1270 | GhA12: 67653347..67655408(-) | 2062 | 4 | 267 | 27 | 29.12 | 8.71 | 31.28 | 77.75 | -0.137 |
GhEXPB2D | Gh_D02G1839 | GhD02: 61451593..61453105(+) | 1513 | 4 | 275 | 34 | 30.25 | 8.98 | 34.08 | 76.22 | -0.242 |
GhEXPB1D | Gh_D08G1516 | GhD08: 48665574..48666719(-) | 1146 | 4 | 273 | NO | 28.61 | 5.03 | 33.99 | 69.71 | -0.015 |
GhEXPB3D | Gh_D12G0941 | GhD12: 34578510..34580023(-) | 1514 | 3 | 180 | NO | 18.92 | 5.03 | 30.62 | 83.44 | 0.032 |
GhEXPB4D | Gh_D12G1393 | GhD12: 43009111..43011170(-) | 2060 | 4 | 267 | 27 | 29.09 | 8.71 | 31.74 | 76.29 | -0.155 |
GhEXLA1A | Gh_A01G1439 | GhA01: 89104034..89105296(+) | 1263 | 5 | 259 | 17 | 28.02 | 8.26 | 34.36 | 83.28 | 0.000 |
GhEXLA2A | Gh_A03G2186 | scaffold739_A03: 13564..15228(-) | 1665 | 4 | 260 | 20 | 28.26 | 8.24 | 35.24 | 82.92 | -0.029 |
GhEXLA3A | Gh_A04G1065 | GhA04: 60713119..60715274(+) | 2156 | 5 | 259 | 18 | 28.10 | 8.59 | 29.34 | 81.39 | -0.019 |
GhEXLA1D | Gh_D01G1679 | GhD01: 52783404..52784664(+) | 1261 | 5 | 259 | 17 | 28.01 | 8.26 | 33.65 | 81.78 | 0.001 |
GhEXLA2D | Gh_D02G2280 | GhD02: 66566105..66567707(+) | 1603 | 4 | 260 | 20 | 28.21 | 8.24 | 33.56 | 81.42 | -0.050 |
GhEXLA3D | Gh_D04G1670 | GhD04: 48878933..48881117(+) | 2185 | 5 | 259 | 18 | 28.13 | 8.59 | 29.82 | 80.27 | -0.027 |
GhEXLB1A | Gh_A03G0046 | GhA03: 643878..645995(+) | 2118 | 4 | 250 | 24 | 27.75 | 6.09 | 28.76 | 74.12 | -0.153 |
GhEXLB2A | Gh_A08G1444 | GhA08: 90327706..90329016(-) | 1311 | 5 | 256 | 24 | 27.85 | 4.81 | 31.94 | 82.62 | -0.099 |
GhEXLB3A | Gh_A08G1477 | GhA08: 90994259..90995358(+) | 1100 | 4 | 249 | 25 | 27.51 | 6.40 | 29.96 | 77.87 | -0.139 |
GhEXLB4A | Gh_A08G1478 | GhA08: 91000911..91001970(+) | 1060 | 4 | 248 | NO | 27.67 | 9.00 | 23.00 | 80.52 | -0.158 |
GhEXLB6A | Gh_A12G1923 | GhA12: 81945320..81946466(-) | 1147 | 5 | 249 | 21 | 27.08 | 4.65 | 35.69 | 84.98 | -0.109 |
GhEXLB1D | Gh_D03G1609 | GhD03: 46020990..46023128(-) | 2139 | 4 | 250 | 24 | 27.69 | 5.64 | 26.36 | 71.76 | -0.174 |
GhEXLB2D | Gh_D08G1739 | GhD08: 53651905..53653218(-) | 1314 | 5 | 256 | 24 | 27.90 | 4.66 | 32.35 | 82.62 | -0.099 |
GhEXLB3D | Gh_D08G1774 | GhD08: 54163295..54164384(+) | 1090 | 4 | 249 | 25 | 27.36 | 6.06 | 31.21 | 79.08 | -0.113 |
GhEXLB4-1D | Gh_D08G1775 | GhD08: 54170354..54171413(+) | 1060 | 4 | 248 | NO | 27.67 | 9.00 | 23.00 | 80.52 | -0.158 |
GhEXLB4-2D | Gh_D08G1776 | GhD08: 54175086..54176147(+) | 1062 | 4 | 248 | 25 | 27.55 | 8.96 | 25.58 | 80.52 | -0.147 |
GhEXLB5D | Gh_D08G1777 | GhD08: 54180358..54181440(+) | 1083 | 4 | 250 | 24 | 27.57 | 6.12 | 28.08 | 77.96 | -0.104 |
GhEXLB6D | Gh_D12G2103 | GhD12: 54022894..54024050(-) | 1157 | 5 | 249 | 21 | 27.12 | 4.65 | 36.35 | 85.38 | -0.098 |
表3
不同植物扩展蛋白及其各亚家族内基因数目"
物种Species | EXPA | EXPB | EXLA | EXLB | 总计Total | 参考文献Reference |
---|---|---|---|---|---|---|
陆地棉G. hirsutum | 46 | 8 | 6 | 12 | 72 | 本研究This study |
亚洲棉G. arboreum | 26 | 4 | 3 | 6 | 39 | 本研究This study |
雷蒙德氏棉G. raimondii | 26 | 4 | 3 | 6 | 39 | [46] |
黄瓜Cucumis sativus | 21 | 3 | 9 | 2 | 35 | [48] |
拟南芥Arabidopsis thaliana | 26 | 6 | 3 | 1 | 36 | [6] |
水稻Oryza sativa | 34 | 19 | 4 | 1 | 58 | [6] |
玉米Zea mays | 36 | 48 | 4 | 0 | 88 | [7] |
大白菜B. rapa (Pekinensis) | 39 | 9 | 2 | 3 | 53 | [50] |
杨树Populus trichocarpa | 27 | 3 | 2 | 4 | 36 | [51] |
葡萄Vitis vinifera | 20 | 4 | 1 | 4 | 29 | [10] |
大豆Glycine max | 49 | 9 | 2 | 5 | 75 | [8] |
苹果Malus×Domestica (Borkh) | 34 | 1 | 2 | 4 | 41 | [52] |
菜豆Phaseolus vulgaris | 25 | 6 | 0 | 5 | 36 | [53] |
苜蓿Medicago truncatula | 16 | 1 | 0 | 1 | 18 | [53] |
小粒碗藓Physcomitrella patens | 27 | 7 | 0 | 0 | 34 | [54] |
共同祖先Last common ancestor | 10—12 | 2 | 1 | 2 | 15—17 | [6] |
表4
陆地棉扩展蛋白基因密码子的相对使用频率"
aa | Cod | Freq | aa | Cod | Freq | aa | Cod | Freq | aa | Cod | Freq | |||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
F | UUU | 0.95 | S | UCU | 1.35 | Y | UAU | 1.06 | C | UGU | 0.99 | |||
UUC | 1.05 | UCC | 0.94 | UAC | 0.94 | UGC | 1.01 | |||||||
L | UUA | 0.77 | UCA | 1.06 | * | UAA | 1.13 | * | UGA | 0.75 | ||||
UUG | 1.54 | UCG | 0.49 | UAG | 1.13 | W | UGG | 1 | ||||||
CUU | 1.22 | P | CCU | 1.69 | H | CAU | 1.26 | R | CGU | 0.54 | ||||
CUC | 1.05 | CCC | 0.73 | CAC | 0.74 | CGC | 0.5 | |||||||
CUA | 0.64 | CCA | 1.24 | Q | CAA | 1.2 | CGA | 0.63 | ||||||
CUG | 0.78 | CCG | 0.34 | CAG | 0.8 | CGG | 0.29 | |||||||
I | AUU | 1.3 | T | ACU | 1.28 | N | AAU | 1.02 | S | AGU | 0.86 | |||
AUC | 1.05 | ACC | 1.14 | AAC | 0.98 | AGC | 1.31 | |||||||
AUA | 0.64 | ACA | 1.33 | K | AAA | 1.11 | R | AGA | 1.8 | |||||
M | AUG | 1 | ACG | 0.26 | AAG | 0.89 | AGG | 2.24 | ||||||
V | GUU | 1.42 | A | GCU | 1.6 | D | GAU | 1.18 | G | GGU | 1.23 | |||
GUC | 0.8 | GCC | 0.91 | GAC | 0.82 | GGC | 0.84 | |||||||
GUA | 0.62 | GCA | 1.19 | E | GAA | 1.18 | GGA | 1.2 | ||||||
GUG | 1.16 | GCG | 0.3 | GAG | 0.82 | GGG | 0.73 |
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