园艺学报 ›› 2022, Vol. 49 ›› Issue (8): 1772-1784.doi: 10.16420/j.issn.0513-353x.2021-0389
        
               		蒋思思1, 袁军1, 周文君1, 钮根花2, 周俊琴1,*(
)
                  
        
        
        
        
    
收稿日期:2021-12-21
									
				
											修回日期:2022-03-08
									
				
									
				
											出版日期:2022-08-25
									
				
											发布日期:2022-09-05
									
			通讯作者:
					周俊琴
											E-mail:zhoujunqin@csuft.edu.cn
												基金资助:
        
               		JIANG Sisi1, YUAN Jun1, ZHOU Wenjun1, NIU Genhua2, ZHOU Junqin1,*(
)
			  
			
			
			
                
        
    
Received:2021-12-21
									
				
											Revised:2022-03-08
									
				
									
				
											Online:2022-08-25
									
				
											Published:2022-09-05
									
			Contact:
					ZHOU Junqin   
											E-mail:zhoujunqin@csuft.edu.cn
												摘要:
利用Illumina平台对薄壳山核桃(Carya illinoinensis)品种‘金华’‘波尼’和1株实生古树单株的叶绿体全基因组进行测序,并对其特征进行分析。3个薄壳山核桃叶绿体基因组均呈典型的四分体结构,大小分别为160 819、162 611和160 762 bp,基因序列高度保守。三者均注释得到124个基因,包括79种蛋白编码基因、38个tRNA和7个rRNA基因;三者有相似的重复序列类型,在其叶绿体基因组序列中分别检测出74、74和73个SSR位点;系统发育分析表明,三者均与喙核桃(Annamocarya sinensis)亲缘关系最为密切。
中图分类号:
蒋思思, 袁军, 周文君, 钮根花, 周俊琴. 薄壳山核桃(Carya illinoinensis)叶绿体基因组及其特征分析[J]. 园艺学报, 2022, 49(8): 1772-1784.
JIANG Sisi, YUAN Jun, ZHOU Wenjun, NIU Genhua, ZHOU Junqin. Complete Chloroplast Genome Sequence and Characteristics Analysis of Carya illinoinensis[J]. Acta Horticulturae Sinica, 2022, 49(8): 1772-1784.
| 品种  Cultivar  |  LSC | SSC | IR | Genome | ||||||
|---|---|---|---|---|---|---|---|---|---|---|
| 长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  | |
| 金华 Jinhua | 90 022 | 33.74 | 55.98 | 18 791 | 29.89 | 11.68 | 26 003 | 42.58 | 16.17 | 160 819 | 
| 波尼 Pawnee | 90 022 | 33.74 | 55.36 | 16 999 | 29.93 | 10.45 | 27 795 | 41.74 | 17.09 | 162 611 | 
| 实生单株  Seedling individual  |  89 964 | 33.75 | 55.96 | 18 792 | 29.88 | 11.69 | 26 003 | 42.58 | 16.17 | 160 762 | 
表1 薄壳山核桃‘波尼’‘金华’和实生单株的叶绿体基因组特征综述
Table 1 Summary of the chloroplast genomes features of Carya illinoinensis‘Pawnee’‘Jinhua’and seedling individual
| 品种  Cultivar  |  LSC | SSC | IR | Genome | ||||||
|---|---|---|---|---|---|---|---|---|---|---|
| 长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  |  G + C/% | 长度/%  Length  |  长度/bp  Length  | |
| 金华 Jinhua | 90 022 | 33.74 | 55.98 | 18 791 | 29.89 | 11.68 | 26 003 | 42.58 | 16.17 | 160 819 | 
| 波尼 Pawnee | 90 022 | 33.74 | 55.36 | 16 999 | 29.93 | 10.45 | 27 795 | 41.74 | 17.09 | 162 611 | 
| 实生单株  Seedling individual  |  89 964 | 33.75 | 55.96 | 18 792 | 29.88 | 11.69 | 26 003 | 42.58 | 16.17 | 160 762 | 
																													图2 薄壳山核桃实生单株叶绿体全基因组图谱 内侧基因顺时针,外侧基因逆时针(Lohse et al.,2013)。
Fig. 2 Gene map of the complete chloroplast genome of Carya illinoinensis seedling individual Genes drawn inside the circle are transcribed clockwise,and those outsides are counter clockwise(Lohse et al.,2013).
| 基因分类  Category for gene  |  基因分组  Group of gene  |  基因名称  Name of gene  | 
|---|---|---|
| 自我复制  Self replication  |  rRNA | rrn16S,rrn23,rrn23S,rrn4.5,rrn4.5S,rrn5,rrn5S | 
| tRNA | trnA-UGC*,trnC-GCA,trnD-GUC,trnE-UUC,trnF-GAA,trnfM-CAU,trnG-GCC*,trnG-UCC,trnH-GUG,trnI-CAU,trnI-GAU*,trnK-UUU*,trnL-CAA,trnL-UAA*,trnL-UAG,trnM-CAU,trnN-GUU,trnP-GGG,trnP-UGG,trnQ-UUG,trnR-ACG,trnR-UCU,trnS-CGA,trnS-GCU,trnS-GGA,trnStop-UUA,trnS-UGA,trnT-AAG,trnT-AGU,trnT-CAG,trnT-GGU,trnT-UAG,trnT-UGU,trnV-AAC,trnV-GAC,trnV-UAC*,trnW-CCA,trnY-GUA | |
| RNA聚合酶亚基  RNA polymerase  |  rpoA,rpoB,rpoC1*,rpoC2 | |
| 核糖体小亚基  Ribosomal protein(SSU)  |  rps2,rps3,rps4,rps7,rps8,rps11,rps12T,*,rps14,rps15,rps16*,rps18,rps19 | |
| 核糖体大亚基  Ribosomal protein(LSU)  |  rpl2*,rpl14,rpl16*,rpl20,rpl22,rpl23,rpl32,rpl33,rpl36 | |
| 光合作用相关  Genes for photo- synthesis  |  ATP 合酶 ATP synthase | atpA,atpB,atpE,atpF*,atpH,atpI | 
| 光和系统Ⅰ Photosystem Ⅰ | psaA,psaB,psaC,psaI,psaJ | |
| 光和系统Ⅱ Photosystem Ⅱ | psbA,psbB,psbC,psbD,psbE,psbF,psbI,psbJ,psbK,psbH,psbL,psbM,psbN,psbT,psbZ,ycf3** | |
| 二磷酸核酮糖羧化酶亚基  Subunit of rubisco  |  rbcL | |
| 细胞色素复合物  Cytochrome b/f complex  |  petA,petB*,petD*,petG,petL,petN | |
| NADH脱氢酶  NADH-dehydrogenase  |  ndhA*,ndhB*,ndhC,ndhD,ndhE,ndhF,ndhG,ndhH,ndhI,ndhJ,ndhK | |
| 其他与未知功能  Other and unknown function  |  accD,ccsA,cemA,clpP**,matK,ycf1,ycf2,ycf4,ycf15a | 
表2 薄壳山核桃叶绿体基因组注释基因列表
Table 2 List of genes annotated in the chloroplast genome of Carya illinoinensis sequenced
| 基因分类  Category for gene  |  基因分组  Group of gene  |  基因名称  Name of gene  | 
|---|---|---|
| 自我复制  Self replication  |  rRNA | rrn16S,rrn23,rrn23S,rrn4.5,rrn4.5S,rrn5,rrn5S | 
| tRNA | trnA-UGC*,trnC-GCA,trnD-GUC,trnE-UUC,trnF-GAA,trnfM-CAU,trnG-GCC*,trnG-UCC,trnH-GUG,trnI-CAU,trnI-GAU*,trnK-UUU*,trnL-CAA,trnL-UAA*,trnL-UAG,trnM-CAU,trnN-GUU,trnP-GGG,trnP-UGG,trnQ-UUG,trnR-ACG,trnR-UCU,trnS-CGA,trnS-GCU,trnS-GGA,trnStop-UUA,trnS-UGA,trnT-AAG,trnT-AGU,trnT-CAG,trnT-GGU,trnT-UAG,trnT-UGU,trnV-AAC,trnV-GAC,trnV-UAC*,trnW-CCA,trnY-GUA | |
| RNA聚合酶亚基  RNA polymerase  |  rpoA,rpoB,rpoC1*,rpoC2 | |
| 核糖体小亚基  Ribosomal protein(SSU)  |  rps2,rps3,rps4,rps7,rps8,rps11,rps12T,*,rps14,rps15,rps16*,rps18,rps19 | |
| 核糖体大亚基  Ribosomal protein(LSU)  |  rpl2*,rpl14,rpl16*,rpl20,rpl22,rpl23,rpl32,rpl33,rpl36 | |
| 光合作用相关  Genes for photo- synthesis  |  ATP 合酶 ATP synthase | atpA,atpB,atpE,atpF*,atpH,atpI | 
| 光和系统Ⅰ Photosystem Ⅰ | psaA,psaB,psaC,psaI,psaJ | |
| 光和系统Ⅱ Photosystem Ⅱ | psbA,psbB,psbC,psbD,psbE,psbF,psbI,psbJ,psbK,psbH,psbL,psbM,psbN,psbT,psbZ,ycf3** | |
| 二磷酸核酮糖羧化酶亚基  Subunit of rubisco  |  rbcL | |
| 细胞色素复合物  Cytochrome b/f complex  |  petA,petB*,petD*,petG,petL,petN | |
| NADH脱氢酶  NADH-dehydrogenase  |  ndhA*,ndhB*,ndhC,ndhD,ndhE,ndhF,ndhG,ndhH,ndhI,ndhJ,ndhK | |
| 其他与未知功能  Other and unknown function  |  accD,ccsA,cemA,clpP**,matK,ycf1,ycf2,ycf4,ycf15a | 
| 基因  Gene  |  正负链  Strand  |  起始 Start | 终止 End | 外显子  Ⅰ/bp ExonⅠ  |  内含子  Ⅰ/bp IntronⅠ  |  外显子  Ⅱ/bp ExonⅡ  |  内含子  Ⅱ/bp Intron Ⅱ  |  外显子  Ⅲ/bp Exon Ⅲ  | ||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | |||||||
| trnK-UUU | - | 1 607 | 1 607 | 1 607 | 4 237 | 4 237 | 4 237 | 37 | 2 559 | 35 | ||
| rps16 | - | 5 086 | 5 086 | 5 086 | 6 257 | 6 257 | 6 257 | 39 | 905 | 228 | ||
| trnG-GCC | + | 10 135 | 10 135 | 10 134 | 10 919 | 10 919 | 10 918 | 23 | 715 | 47 | ||
| atpF | - | 13 185 | 13 185 | 13 184 | 14 490 | 14 490 | 14 489 | 145 | 751 | 410 | ||
| rpoC1 | - | 22 810 | 22 810 | 22 811 | 25 681 | 25 681 | 25 683 | 432 | 814 | 1 626 | ||
| ycf3 | - | 47 097 | 47 097 | 47 034 | 49 116 | 49 116 | 49 053 | 124 | 720 | 230 | 793 | 153 | 
| trnL-UAA | + | 52 205 | 52 205 | 52 142 | 52 815 | 52 815 | 52 752 | 37 | 524 | 50 | ||
| trnV-UAC | - | 56 413 | 56 413 | 56 350 | 57 102 | 57 102 | 57 039 | 38 | 615 | 37 | ||
| clpP | - | 75 722 | 75 722 | 75 665 | 77 773 | 77 773 | 77 715 | 71 | 847 | 294 | 614 | 226 | 
| petB | + | 80 745 | 80 745 | 80 687 | 82 207 | 82 207 | 82 149 | 6 | 815 | 642 | ||
| petD | + | 82 418 | 82 418 | 82 360 | 83 526 | 83 526 | 83 468 | 9 | 626 | 474 | ||
| rpl16 | - | 87 056 | 87 056 | 86 998 | 88 382 | 88 382 | 88 324 | 9 | 919 | 399 | ||
| rpl2 | - | 90 103 | 90 103 | 90 045 | 91 625 | 91 625 | 91 567 | 391 | 698 | 434 | ||
| ndhB | - | 100 446 | 100 446 | 100 388 | 102 670 | 102 670 | 102 612 | 775 | 686 | 764 | ||
| trnI-GAU | + | 108 073 | 108 073 | 108 015 | 109 099 | 109 099 | 109 041 | 37 | 955 | 35 | ||
| trnA-UGC | + | 109 164 | 109 164 | 109 106 | 110 043 | 110 043 | 109 985 | 38 | 807 | 35 | ||
| ndhA | - | 125 970 | 127 762 | 125 912 | 128 253 | 130 045 | 128 195 | 553 | 1 192 | 539 | ||
| trnA-UGC | - | 140 799 | 142 591 | 140 742 | 141 678 | 143 470 | 141 621 | 38 | 807 | 35 | ||
| trnI-GAU | - | 141 743 | 143 535 | 141 686 | 142 769 | 144 561 | 142 712 | 37 | 955 | 35 | ||
| ndhB | + | 148 172 | 149 964 | 148 115 | 150 396 | 152 188 | 150 339 | 775 | 686 | 764 | ||
| rpl2 | + | 159 217 | 161 009 | 159 160 | 160 739 | 162 531 | 160 682 | 391 | 698 | 434 | ||
表3 薄壳山核桃‘金华’‘波尼’和实生单株叶绿体基因组中内含子和外显子长度
Table 3 Intron and exon length in the chloroplast genome of Carya illinoinensis‘Jinhua’‘Pawnee’and seedling individual
| 基因  Gene  |  正负链  Strand  |  起始 Start | 终止 End | 外显子  Ⅰ/bp ExonⅠ  |  内含子  Ⅰ/bp IntronⅠ  |  外显子  Ⅱ/bp ExonⅡ  |  内含子  Ⅱ/bp Intron Ⅱ  |  外显子  Ⅲ/bp Exon Ⅲ  | ||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | |||||||
| trnK-UUU | - | 1 607 | 1 607 | 1 607 | 4 237 | 4 237 | 4 237 | 37 | 2 559 | 35 | ||
| rps16 | - | 5 086 | 5 086 | 5 086 | 6 257 | 6 257 | 6 257 | 39 | 905 | 228 | ||
| trnG-GCC | + | 10 135 | 10 135 | 10 134 | 10 919 | 10 919 | 10 918 | 23 | 715 | 47 | ||
| atpF | - | 13 185 | 13 185 | 13 184 | 14 490 | 14 490 | 14 489 | 145 | 751 | 410 | ||
| rpoC1 | - | 22 810 | 22 810 | 22 811 | 25 681 | 25 681 | 25 683 | 432 | 814 | 1 626 | ||
| ycf3 | - | 47 097 | 47 097 | 47 034 | 49 116 | 49 116 | 49 053 | 124 | 720 | 230 | 793 | 153 | 
| trnL-UAA | + | 52 205 | 52 205 | 52 142 | 52 815 | 52 815 | 52 752 | 37 | 524 | 50 | ||
| trnV-UAC | - | 56 413 | 56 413 | 56 350 | 57 102 | 57 102 | 57 039 | 38 | 615 | 37 | ||
| clpP | - | 75 722 | 75 722 | 75 665 | 77 773 | 77 773 | 77 715 | 71 | 847 | 294 | 614 | 226 | 
| petB | + | 80 745 | 80 745 | 80 687 | 82 207 | 82 207 | 82 149 | 6 | 815 | 642 | ||
| petD | + | 82 418 | 82 418 | 82 360 | 83 526 | 83 526 | 83 468 | 9 | 626 | 474 | ||
| rpl16 | - | 87 056 | 87 056 | 86 998 | 88 382 | 88 382 | 88 324 | 9 | 919 | 399 | ||
| rpl2 | - | 90 103 | 90 103 | 90 045 | 91 625 | 91 625 | 91 567 | 391 | 698 | 434 | ||
| ndhB | - | 100 446 | 100 446 | 100 388 | 102 670 | 102 670 | 102 612 | 775 | 686 | 764 | ||
| trnI-GAU | + | 108 073 | 108 073 | 108 015 | 109 099 | 109 099 | 109 041 | 37 | 955 | 35 | ||
| trnA-UGC | + | 109 164 | 109 164 | 109 106 | 110 043 | 110 043 | 109 985 | 38 | 807 | 35 | ||
| ndhA | - | 125 970 | 127 762 | 125 912 | 128 253 | 130 045 | 128 195 | 553 | 1 192 | 539 | ||
| trnA-UGC | - | 140 799 | 142 591 | 140 742 | 141 678 | 143 470 | 141 621 | 38 | 807 | 35 | ||
| trnI-GAU | - | 141 743 | 143 535 | 141 686 | 142 769 | 144 561 | 142 712 | 37 | 955 | 35 | ||
| ndhB | + | 148 172 | 149 964 | 148 115 | 150 396 | 152 188 | 150 339 | 775 | 686 | 764 | ||
| rpl2 | + | 159 217 | 161 009 | 159 160 | 160 739 | 162 531 | 160 682 | 391 | 698 | 434 | ||
| 密码子Codon | 氨基酸  Amino acid  |  频率/%  Frequency  |  数量  Number  |  RSCU | ||||||
|---|---|---|---|---|---|---|---|---|---|---|
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株 Seedling individual | 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | ||
| GCA | A | 1.454 | 1.426 | 1.454 | 715 | 729 | 715 | 1.130 | 1.140 | 1.130 | 
| GCC | A | 0.679 | 0.665 | 0.679 | 334 | 340 | 334 | 0.530 | 0.530 | 0.530 | 
| GCG | A | 0.596 | 0.585 | 0.592 | 293 | 299 | 291 | 0.460 | 0.470 | 0.460 | 
| GCT | A | 2.408 | 2.348 | 2.408 | 1 184 | 1 200 | 1 184 | 1.870 | 1.870 | 1.880 | 
| TGC | C | 0.334 | 0.333 | 0.334 | 164 | 170 | 164 | 0.560 | 0.560 | 0.560 | 
| TGT | C | 0.860 | 0.855 | 0.860 | 423 | 437 | 423 | 1.440 | 1.440 | 1.440 | 
| GAC | D | 0.810 | 0.802 | 0.810 | 398 | 410 | 398 | 0.400 | 0.390 | 0.400 | 
| GAT | D | 3.259 | 3.272 | 3.259 | 1 602 | 1 672 | 1 602 | 1.600 | 1.610 | 1.600 | 
| GAA | E | 3.897 | 3.996 | 3.897 | 1 916 | 2 042 | 1 916 | 1.490 | 1.500 | 1.490 | 
| GAG | E | 1.328 | 1.336 | 1.332 | 653 | 683 | 655 | 0.510 | 0.500 | 0.510 | 
| TTC | F | 2.018 | 2.015 | 2.018 | 992 | 1 030 | 992 | 0.690 | 0.680 | 0.690 | 
| TTT | F | 3.794 | 3.872 | 3.794 | 1 865 | 1 979 | 1 865 | 1.310 | 1.320 | 1.310 | 
| GGA | G | 2.532 | 2.499 | 2.532 | 1 245 | 1 277 | 1 245 | 1.590 | 1.600 | 1.590 | 
| GGC | G | 0.712 | 0.701 | 0.712 | 350 | 358 | 350 | 0.450 | 0.450 | 0.450 | 
| GGG | G | 0.970 | 0.961 | 0.970 | 477 | 491 | 477 | 0.610 | 0.620 | 0.610 | 
| GGT | G | 2.140 | 2.090 | 2.140 | 1 052 | 1 068 | 1 052 | 1.350 | 1.340 | 1.350 | 
| CAC | H | 0.547 | 0.546 | 0.547 | 269 | 279 | 269 | 0.450 | 0.450 | 0.450 | 
| CAT | H | 1.906 | 1.869 | 1.906 | 937 | 955 | 937 | 1.550 | 1.550 | 1.550 | 
| ATA | I | 2.888 | 2.947 | 2.893 | 1 420 | 1 506 | 1 422 | 0.990 | 1.000 | 0.990 | 
| ATC | I | 1.621 | 1.622 | 1.621 | 797 | 829 | 797 | 0.550 | 0.550 | 0.550 | 
| ATT | I | 4.274 | 4.283 | 4.274 | 2 101 | 2 189 | 2 101 | 1.460 | 1.450 | 1.460 | 
| AAA | K | 3.995 | 4.133 | 3.995 | 1 964 | 2 112 | 1 964 | 1.450 | 1.450 | 1.450 | 
| AAG | K | 1.511 | 1.552 | 1.507 | 743 | 793 | 741 | 0.550 | 0.550 | 0.550 | 
| CTA | L | 1.336 | 1.333 | 1.332 | 657 | 681 | 655 | 0.760 | 0.760 | 0.760 | 
| CTC | L | 0.755 | 0.757 | 0.755 | 371 | 387 | 371 | 0.430 | 0.430 | 0.430 | 
| CTG | L | 0.675 | 0.681 | 0.675 | 332 | 348 | 332 | 0.390 | 0.390 | 0.390 | 
| CTT | L | 2.258 | 2.227 | 2.258 | 1 110 | 1 138 | 1 110 | 1.290 | 1.280 | 1.290 | 
| TTA | L | 3.242 | 3.260 | 3.242 | 1 594 | 1 666 | 1 594 | 1.850 | 1.870 | 1.850 | 
| TTG | L | 2.246 | 2.199 | 2.246 | 1 104 | 1 124 | 1 104 | 1.280 | 1.260 | 1.280 | 
| ATG | M | 2.244 | 2.225 | 2.244 | 1 103 | 1 137 | 1 103 | 1.000 | 1.000 | 1.000 | 
| AAC | N | 1.153 | 1.156 | 1.153 | 567 | 591 | 567 | 0.470 | 0.460 | 0.470 | 
| AAT | N | 3.794 | 3.837 | 3.802 | 1 865 | 1 961 | 1 869 | 1.530 | 1.540 | 1.530 | 
| 密码子Codon | 氨基酸  Amino acid  |  频率/%  Frequency  |  数量  Number  |  RSCU | ||||||
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株 Seedling individual | 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | ||
| CCA | P | 1.168 | 1.155 | 1.168 | 574 | 590 | 574 | 1.140 | 1.130 | 1.140 | 
| CCC | P | 0.816 | 0.816 | 0.816 | 401 | 417 | 401 | 0.800 | 0.800 | 0.800 | 
| CCG | P | 0.633 | 0.640 | 0.633 | 311 | 327 | 311 | 0.620 | 0.630 | 0.620 | 
| CCT | P | 1.479 | 1.477 | 1.479 | 727 | 755 | 727 | 1.440 | 1.450 | 1.440 | 
| CAA | Q | 2.805 | 2.820 | 2.805 | 1 379 | 1 441 | 1 379 | 1.550 | 1.560 | 1.550 | 
| CAG | Q | 0.818 | 0.806 | 0.818 | 402 | 412 | 402 | 0.450 | 0.440 | 0.450 | 
| AGA | R | 1.827 | 1.828 | 1.827 | 898 | 934 | 898 | 1.810 | 1.820 | 1.810 | 
| AGG | R | 0.716 | 0.704 | 0.716 | 352 | 360 | 352 | 0.710 | 0.700 | 0.710 | 
| CGA | R | 1.367 | 1.374 | 1.367 | 672 | 702 | 672 | 1.350 | 1.370 | 1.350 | 
| CGC | R | 0.393 | 0.389 | 0.393 | 193 | 199 | 193 | 0.390 | 0.390 | 0.390 | 
| CGG | R | 0.472 | 0.458 | 0.472 | 232 | 234 | 232 | 0.470 | 0.460 | 0.470 | 
| CGT | R | 1.288 | 1.274 | 1.288 | 633 | 651 | 633 | 1.270 | 1.270 | 1.270 | 
| AGC | S | 0.535 | 0.538 | 0.535 | 263 | 275 | 263 | 0.420 | 0.420 | 0.420 | 
| AGT | S | 1.530 | 1.511 | 1.526 | 752 | 772 | 750 | 1.200 | 1.190 | 1.200 | 
| TCA | S | 1.526 | 1.542 | 1.526 | 750 | 788 | 750 | 1.200 | 1.220 | 1.200 | 
| TCC | S | 1.202 | 1.192 | 1.202 | 591 | 609 | 591 | 0.940 | 0.940 | 0.940 | 
| TCG | S | 0.738 | 0.726 | 0.738 | 363 | 371 | 363 | 0.580 | 0.570 | 0.580 | 
| TCT | S | 2.126 | 2.107 | 2.126 | 1 045 | 1 077 | 1 045 | 1.670 | 1.660 | 1.670 | 
| ACA | T | 1.471 | 1.493 | 1.471 | 723 | 763 | 723 | 1.220 | 1.230 | 1.220 | 
| ACC | T | 0.828 | 0.828 | 0.828 | 407 | 423 | 407 | 0.690 | 0.680 | 0.690 | 
| ACG | T | 0.531 | 0.530 | 0.531 | 261 | 271 | 261 | 0.440 | 0.440 | 0.440 | 
| ACT | T | 1.991 | 1.998 | 1.991 | 979 | 1 021 | 979 | 1.650 | 1.650 | 1.650 | 
| GTA | V | 1.983 | 1.970 | 1.983 | 975 | 1 007 | 975 | 1.530 | 1.530 | 1.530 | 
| GTC | V | 0.631 | 0.638 | 0.631 | 310 | 326 | 310 | 0.490 | 0.500 | 0.490 | 
| GTG | V | 0.677 | 0.663 | 0.677 | 333 | 339 | 333 | 0.520 | 0.520 | 0.520 | 
| GTT | V | 1.906 | 1.869 | 1.906 | 937 | 955 | 937 | 1.470 | 1.450 | 1.470 | 
| TGG | W | 1.855 | 1.847 | 1.855 | 912 | 944 | 912 | 1.000 | 1.000 | 1.000 | 
| TAC | Y | 0.767 | 0.753 | 0.767 | 377 | 385 | 377 | 0.410 | 0.400 | 0.410 | 
| TAT | Y | 3.015 | 3.021 | 3.015 | 1 482 | 1 544 | 1 482 | 1.590 | 1.600 | 1.590 | 
| TAA | * | 0.283 | 0.276 | 0.283 | 139 | 141 | 139 | 1.260 | 1.270 | 1.260 | 
| TAG | * | 0.205 | 0.198 | 0.205 | 101 | 101 | 101 | 0.920 | 0.910 | 0.920 | 
| TGA | * | 0.185 | 0.178 | 0.185 | 91 | 91 | 91 | 0.830 | 0.820 | 0.830 | 
表4 薄壳山核桃‘金华’‘波尼’和实生单株叶绿体基因相对同义密码子使用度
Table 4 Relative synonymous codon usage in chloroplast genes of Carya illinoinensis‘Jinhua’‘Pawnee’and Seedling individual
| 密码子Codon | 氨基酸  Amino acid  |  频率/%  Frequency  |  数量  Number  |  RSCU | ||||||
|---|---|---|---|---|---|---|---|---|---|---|
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株 Seedling individual | 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | ||
| GCA | A | 1.454 | 1.426 | 1.454 | 715 | 729 | 715 | 1.130 | 1.140 | 1.130 | 
| GCC | A | 0.679 | 0.665 | 0.679 | 334 | 340 | 334 | 0.530 | 0.530 | 0.530 | 
| GCG | A | 0.596 | 0.585 | 0.592 | 293 | 299 | 291 | 0.460 | 0.470 | 0.460 | 
| GCT | A | 2.408 | 2.348 | 2.408 | 1 184 | 1 200 | 1 184 | 1.870 | 1.870 | 1.880 | 
| TGC | C | 0.334 | 0.333 | 0.334 | 164 | 170 | 164 | 0.560 | 0.560 | 0.560 | 
| TGT | C | 0.860 | 0.855 | 0.860 | 423 | 437 | 423 | 1.440 | 1.440 | 1.440 | 
| GAC | D | 0.810 | 0.802 | 0.810 | 398 | 410 | 398 | 0.400 | 0.390 | 0.400 | 
| GAT | D | 3.259 | 3.272 | 3.259 | 1 602 | 1 672 | 1 602 | 1.600 | 1.610 | 1.600 | 
| GAA | E | 3.897 | 3.996 | 3.897 | 1 916 | 2 042 | 1 916 | 1.490 | 1.500 | 1.490 | 
| GAG | E | 1.328 | 1.336 | 1.332 | 653 | 683 | 655 | 0.510 | 0.500 | 0.510 | 
| TTC | F | 2.018 | 2.015 | 2.018 | 992 | 1 030 | 992 | 0.690 | 0.680 | 0.690 | 
| TTT | F | 3.794 | 3.872 | 3.794 | 1 865 | 1 979 | 1 865 | 1.310 | 1.320 | 1.310 | 
| GGA | G | 2.532 | 2.499 | 2.532 | 1 245 | 1 277 | 1 245 | 1.590 | 1.600 | 1.590 | 
| GGC | G | 0.712 | 0.701 | 0.712 | 350 | 358 | 350 | 0.450 | 0.450 | 0.450 | 
| GGG | G | 0.970 | 0.961 | 0.970 | 477 | 491 | 477 | 0.610 | 0.620 | 0.610 | 
| GGT | G | 2.140 | 2.090 | 2.140 | 1 052 | 1 068 | 1 052 | 1.350 | 1.340 | 1.350 | 
| CAC | H | 0.547 | 0.546 | 0.547 | 269 | 279 | 269 | 0.450 | 0.450 | 0.450 | 
| CAT | H | 1.906 | 1.869 | 1.906 | 937 | 955 | 937 | 1.550 | 1.550 | 1.550 | 
| ATA | I | 2.888 | 2.947 | 2.893 | 1 420 | 1 506 | 1 422 | 0.990 | 1.000 | 0.990 | 
| ATC | I | 1.621 | 1.622 | 1.621 | 797 | 829 | 797 | 0.550 | 0.550 | 0.550 | 
| ATT | I | 4.274 | 4.283 | 4.274 | 2 101 | 2 189 | 2 101 | 1.460 | 1.450 | 1.460 | 
| AAA | K | 3.995 | 4.133 | 3.995 | 1 964 | 2 112 | 1 964 | 1.450 | 1.450 | 1.450 | 
| AAG | K | 1.511 | 1.552 | 1.507 | 743 | 793 | 741 | 0.550 | 0.550 | 0.550 | 
| CTA | L | 1.336 | 1.333 | 1.332 | 657 | 681 | 655 | 0.760 | 0.760 | 0.760 | 
| CTC | L | 0.755 | 0.757 | 0.755 | 371 | 387 | 371 | 0.430 | 0.430 | 0.430 | 
| CTG | L | 0.675 | 0.681 | 0.675 | 332 | 348 | 332 | 0.390 | 0.390 | 0.390 | 
| CTT | L | 2.258 | 2.227 | 2.258 | 1 110 | 1 138 | 1 110 | 1.290 | 1.280 | 1.290 | 
| TTA | L | 3.242 | 3.260 | 3.242 | 1 594 | 1 666 | 1 594 | 1.850 | 1.870 | 1.850 | 
| TTG | L | 2.246 | 2.199 | 2.246 | 1 104 | 1 124 | 1 104 | 1.280 | 1.260 | 1.280 | 
| ATG | M | 2.244 | 2.225 | 2.244 | 1 103 | 1 137 | 1 103 | 1.000 | 1.000 | 1.000 | 
| AAC | N | 1.153 | 1.156 | 1.153 | 567 | 591 | 567 | 0.470 | 0.460 | 0.470 | 
| AAT | N | 3.794 | 3.837 | 3.802 | 1 865 | 1 961 | 1 869 | 1.530 | 1.540 | 1.530 | 
| 密码子Codon | 氨基酸  Amino acid  |  频率/%  Frequency  |  数量  Number  |  RSCU | ||||||
| 金华  Jinhua  |  波尼  Pawnee  |  实生单株 Seedling individual | 金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  |  金华  Jinhua  |  波尼  Pawnee  |  实生单株  Seedling individual  | ||
| CCA | P | 1.168 | 1.155 | 1.168 | 574 | 590 | 574 | 1.140 | 1.130 | 1.140 | 
| CCC | P | 0.816 | 0.816 | 0.816 | 401 | 417 | 401 | 0.800 | 0.800 | 0.800 | 
| CCG | P | 0.633 | 0.640 | 0.633 | 311 | 327 | 311 | 0.620 | 0.630 | 0.620 | 
| CCT | P | 1.479 | 1.477 | 1.479 | 727 | 755 | 727 | 1.440 | 1.450 | 1.440 | 
| CAA | Q | 2.805 | 2.820 | 2.805 | 1 379 | 1 441 | 1 379 | 1.550 | 1.560 | 1.550 | 
| CAG | Q | 0.818 | 0.806 | 0.818 | 402 | 412 | 402 | 0.450 | 0.440 | 0.450 | 
| AGA | R | 1.827 | 1.828 | 1.827 | 898 | 934 | 898 | 1.810 | 1.820 | 1.810 | 
| AGG | R | 0.716 | 0.704 | 0.716 | 352 | 360 | 352 | 0.710 | 0.700 | 0.710 | 
| CGA | R | 1.367 | 1.374 | 1.367 | 672 | 702 | 672 | 1.350 | 1.370 | 1.350 | 
| CGC | R | 0.393 | 0.389 | 0.393 | 193 | 199 | 193 | 0.390 | 0.390 | 0.390 | 
| CGG | R | 0.472 | 0.458 | 0.472 | 232 | 234 | 232 | 0.470 | 0.460 | 0.470 | 
| CGT | R | 1.288 | 1.274 | 1.288 | 633 | 651 | 633 | 1.270 | 1.270 | 1.270 | 
| AGC | S | 0.535 | 0.538 | 0.535 | 263 | 275 | 263 | 0.420 | 0.420 | 0.420 | 
| AGT | S | 1.530 | 1.511 | 1.526 | 752 | 772 | 750 | 1.200 | 1.190 | 1.200 | 
| TCA | S | 1.526 | 1.542 | 1.526 | 750 | 788 | 750 | 1.200 | 1.220 | 1.200 | 
| TCC | S | 1.202 | 1.192 | 1.202 | 591 | 609 | 591 | 0.940 | 0.940 | 0.940 | 
| TCG | S | 0.738 | 0.726 | 0.738 | 363 | 371 | 363 | 0.580 | 0.570 | 0.580 | 
| TCT | S | 2.126 | 2.107 | 2.126 | 1 045 | 1 077 | 1 045 | 1.670 | 1.660 | 1.670 | 
| ACA | T | 1.471 | 1.493 | 1.471 | 723 | 763 | 723 | 1.220 | 1.230 | 1.220 | 
| ACC | T | 0.828 | 0.828 | 0.828 | 407 | 423 | 407 | 0.690 | 0.680 | 0.690 | 
| ACG | T | 0.531 | 0.530 | 0.531 | 261 | 271 | 261 | 0.440 | 0.440 | 0.440 | 
| ACT | T | 1.991 | 1.998 | 1.991 | 979 | 1 021 | 979 | 1.650 | 1.650 | 1.650 | 
| GTA | V | 1.983 | 1.970 | 1.983 | 975 | 1 007 | 975 | 1.530 | 1.530 | 1.530 | 
| GTC | V | 0.631 | 0.638 | 0.631 | 310 | 326 | 310 | 0.490 | 0.500 | 0.490 | 
| GTG | V | 0.677 | 0.663 | 0.677 | 333 | 339 | 333 | 0.520 | 0.520 | 0.520 | 
| GTT | V | 1.906 | 1.869 | 1.906 | 937 | 955 | 937 | 1.470 | 1.450 | 1.470 | 
| TGG | W | 1.855 | 1.847 | 1.855 | 912 | 944 | 912 | 1.000 | 1.000 | 1.000 | 
| TAC | Y | 0.767 | 0.753 | 0.767 | 377 | 385 | 377 | 0.410 | 0.400 | 0.410 | 
| TAT | Y | 3.015 | 3.021 | 3.015 | 1 482 | 1 544 | 1 482 | 1.590 | 1.600 | 1.590 | 
| TAA | * | 0.283 | 0.276 | 0.283 | 139 | 141 | 139 | 1.260 | 1.270 | 1.260 | 
| TAG | * | 0.205 | 0.198 | 0.205 | 101 | 101 | 101 | 0.920 | 0.910 | 0.920 | 
| TGA | * | 0.185 | 0.178 | 0.185 | 91 | 91 | 91 | 0.830 | 0.820 | 0.830 | 
																													图3 薄壳山核桃‘金华’‘波尼’和实生单株叶绿体基因组LSC、SSC和IR区边界的比较
Fig. 3 Comparison of LSC,SSC and IR boundary of chloroplast genomes of Carya illinoinensis‘Jinhua’‘Pawnee’and seedling individual
																													图4 基于叶绿体基因组串联序列构建18个物种的最大似然(ML)系统进化树
Fig. 4 Maximum likelihood(ML)phylogenetic tree construction including 18 species based on concatenated sequences from all chloroplast genomes
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