新疆野苹果与‘元帅’‘金冠’的叶绿体基因组比对研究

doi:10.16420/j.issn.0513-353x.2021-0378

摘要/Abstract

摘要：

新疆野苹果（Malus sieversii）被认为是现代栽培苹果的祖先物种。为有效保护苹果遗传资源,揭示新疆野苹果和栽培苹果叶绿体基因组的差异,验证现代栽培苹果是否起源于新疆野苹果。对新疆野苹果和栽培苹果‘元帅’和‘金冠’的叶绿体全基因组进行了测序、组装和注释,获得了其叶绿体基因组物理图谱,并进行了比较基因组分析和系统发育分析。结果表明,新疆野苹果和两个栽培苹果品种的叶绿体基因组均为四段式结构,其基因组总长度在160 068 ~ 160 288 bp之间;共注释出131个基因,其中蛋白编码基因86个,tRNA基因37个,rRNA基因8个;新疆野苹果与栽培苹果的碱基组成也基本相似,AT/GC含量基本相同;新疆野苹果鉴定出43个重复序列和64个简单重复序列位点,栽培品种‘元帅’鉴定出49个重复序列和61个简单重复序列位点,‘金冠’鉴定出43个重复序列和57个简单重复序列位点。系统发育分析结果表明：基于全序列构建的贝叶斯进化树在各个节点具有很高的支持率,苹果属的所有物种形成一个单系群（支持率100%）,新疆野苹果和两个栽培苹果品种聚在一起,形成一个小分支（支持率100%）,且新疆野苹果位于该小分支的基部,验证了新疆野苹果是栽培苹果的祖先物种的推断。

关键词: 新疆野苹果, 叶绿体基因组, 差异分析, 系统发育

Abstract:

The wild apple tree Malus sieversii（Rosaceae）is considered to be the ancestor of modern cultivated apple trees. To effectively conserve apple genetic resources,it would be important to elucidate differences of chloroplast genomes between M. sieversii and cultivated apples,as well as verify whether modern cultivated apples originated from M. sieversii. In this study,we used the next-generation sequencing technology to complete the whole-genome sequencing,assembly,and annotation of M. sieversii as well as two domesticated apple trees‘Red Delicious’and‘Golden Delicious’. We successfully mapped the chloroplast genomes of M. sieversii and the two domesticated apple trees,and subsequently constructed the Malus phylogenetic tree at the level of the chloroplast genome. Through a series of comparative analyses,We obtained the following main results：the chloroplast genome of M. sieversii exhibites a typical quadripartite structure,and the complete genomes ranged between 160 068 and 160 288 bp in length. A total of 131 genes were annotated in M. sieversii and the two domesticated apple trees,including 86 protein-coding,37 tRNA,and eight rRNA genes. The base composition of the chloroplast genomes of M. sieversii and the two domesticated apples varieties were similar. A total of 43,49,and 43 repetitive sequence and 64,61 and 57 simple sequence repeat sites were identified in M. sieversii,‘Red Delicious’and‘Golden Delicious’,respectively. For the simple sequence repeats,the main repeat types were single A or T base repeats. Finally,phylogeny study showed that all the Malus species formed a single monophyletic group（BP = 100）. M. sieversii and the two cultivated varieties were grouped in a subclade with strong support（BP = 100）,supporting the hypothesis that M. sieversii is the putative ancestor of domesticated apples.

Key words: Malus sieversii, chloroplastgenome, variation analysis, phylogeny

中图分类号:

S661.1

丁志杰, 包金波, 柔鲜古丽, 朱甜甜, 李雪丽, 苗浩宇, 田新民. 新疆野苹果与‘元帅’‘金冠’的叶绿体基因组比对研究[J]. 园艺学报, 2022, 49(9): 1977-1990.

DING Zhijie, BAO Jinbo, ROUXIAN Guli, ZHU Tiantian, LI Xueli, MIAO Haoyu, TIAN Xinmin. Comparative Chloroplast Genome Study of Mallus servisii‘Red Delicious’and‘Golden Delicious’[J]. Acta Horticulturae Sinica, 2022, 49(9): 1977-1990.

图/表 10

参考文献 43

[1]	Behura W. 2013. Codon usage bias:causative factors,quantification methods and genome‐wide patterns:with emphasis on insect genomes Biological Reviews, 88 (1):49-61.
[2]	Bianco L, Cestaro A, Sargent D J, Banchi E, Derdak S, Salvi S, Jansen J. 2014. Development and validation of a 20K single nucleotide polymorphism (SNP) whole genome genotyping array for apple(Malus domestica Borkh.). Public Library of Science ONE, 9 (10):e110377.
[3]	Cahoon A B, Sharpe R M, Mysayphonh C, Thompson E J, Ward A D, Lin A H. 2010. The complete chloroplast genome of tall fescue(Lolium arundinaceum;Poaceae)and comparison of whole plastomes from the family Poaceae. American Journal of Botany, 97 (1):49-58. doi: 10.3732/ajb.0900008 pmid: 21622366
[4]	Chagne D, Krieger C, Rassam. 2012. QTL and candidate gene mapping for polyphenolic composition in apple fruit. BMC Plant Biology, 12:12.
[5]	Daccord N, Celton JM, Linsmith G, Becker C. 2017. High-quality de novo assembly of the apple genome and methylome dynamics of early fruit development. Nature Genetics, 49 (7):1099-1106. doi: 10.1038/ng.3886 URL
[6]	Depamphilis C W, Palmer J D. 1990. Loss of photosynthetic and chlororespiratory genes from the plastid genome of a parasitic flowering plant. Nature, 348 (6299):337-339. doi: 10.1038/348337a0 URL
[7]	Duan N B, Bai Y, Sun H H, Wang N, Ma Y M, Li M J, Wang X. 2017. Genome resequencing reveals the history of apple and supports a two-stage model for fruit enlargement. Nature Communications, 8 (1):17-20. doi: 10.1038/s41467-017-00019-3 URL
[8]	Fan W B, Ying W, Jiao Y, Shahzad K, Li Z H. 2018. Comparative chloroplast genomics of dipsacales species:insights into sequence variation,adaptive evolution,and phylogenetic relationships. Frontiers in Plant Science, 9:689-702. doi: 10.3389/fpls.2018.00689 URL
[9]	Flora of China Editorial Committee,CAS. 1974. Flora of China. Vol. 36. Beijing:Science Press:372-402. (in Chinese)
	中科院中国植物志编辑委员会. 1974. 中国植物志. 36卷. 北京:科学出版社:372-402.
[10]	Gao Q, Yue G D, Li W Q, Wang J Y, Xu J H. 2012. Recent progress using high-throughput sequencing technologies in plant molecular breeding. Journal of Integrative Plant Biology, 54:215-227. doi: 10.1111/j.1744-7909.2012.01115.x
[11]	Gao Yuan, Wang Dajiang, Wang Kun, Cong Peihua, Li Lianwen, Piao Jicheng. 2020. Analysis of genetic diversity of apple germplasms of Malus using SLAF-seq technology. Acta Horticulturae Sinica, 47 (10):1869-1882. (in Chinese)
	高源, 王大江, 王昆, 丛佩华, 李连文, 朴继成. 2020. 苹果属植物种质多样性的SLAF-seq分析. 园艺学报, 47 (10):1869-1882.
[12]	Goulding S E, Olmstead R G. 1996. Ebb and flow of the chloroplast inverted repeat. Molecular and General Genetics, 252 (2):195-206. doi: 10.1007/BF02173220 URL
[13]	Huang D I, Cronk Q C B. 2015. Plann:A command-line application for annotating plastome sequences. Appl Plant Sci, 3 (8):1500026. doi: 10.3732/apps.1500026 URL
[14]	Katoh K, Standley M. 2013. MAFFT Multiple sequence alignment software version7:improvements in performance and usability. Molecular Biology and Evolution, 30 (4):772-780. doi: 10.1093/molbev/mst010 URL
[15]	Khan MA, Olsen KM, Sovero V, Kushad M M, Schuyler K. 2014. Fruit quality traits have played criticalRoles in domestication of the apple. Plant Genome, 7 (3):doi-10.
[16]	Kumar S, Chagne D, Bink MC, Volz R K, Carlisle C. 2012. Genomic selection for fruit quality traits in apple(Malus domestica Borkh.). Public Library of Science ONE, 7:e36674.
[17]	Kumar S, Garrick D J, Bink M C, Whitworth C, Volz R K. 2013. Novel genomic approaches unravel genetic architecture of complex traits in apple. BMC Genomics, 14:393. doi: 10.1186/1471-2164-14-393 pmid: 23758946
[18]	Kumar S, Raulier P, Chagne D, Whitworth C. 2014. Molecularlevel and trait-level differentiation between the cultivated apple(Malus domestica Borkh.)and its main progenitor Malus sieversii. Plant Genetic Resources, 12:330-340. doi: 10.1017/S1479262114000136 URL
[19]	Kumar S, Rowan D, Hunt M, Chagne D, Whitworth C. 2015. Genome-wide scans reveal genetic architecture of apple flavor volatiles. Molecular Breeding, 35:118. doi: 10.1007/s11032-015-0312-7 URL
[20]	Kurtz S, Choudhuri J V, Ohlebusch E, Schleiermacher C, Stoye J, Giegerich R. 2001. REPuter:the manifold applications of repeat analysis on a genomic scale. Nucleic Acids Research, 29 (22):4633-4642. pmid: 11713313
[21]	Lee S B, Kaittanis C, Jansen R K, Hostetler J B, Tallon L J, Town C D, Daniellet H. 2006. The complete chloroplast genome sequence of Gossypium hirsutum:organization and phylogenetic relationships to other angiosperms. BMC Genomics, 7 (1):61-70. doi: 10.1186/1471-2164-7-61 URL
[22]	Leforestier D, Ravon E, Muranty H, Cornille A, Lemaire C, Giraud T. 2015. Genomic basis of the differences between cider and dessert apple varieties. Evolutionary Applications, 8:650-661. doi: 10.1111/eva.12270 pmid: 26240603
[23]	Li Qian, Guo Qiqiang, Gao Chao, Li Huie. 2020. Characterization of complete chloroplast genome of Camellia weiningensis in Weining,Guizhou Province. Acta Horticulturae Sinica, 47 (4):779-787. (in Chinese)
	李倩, 郭其强, 高超, 李慧娥. 2020. 贵州威宁红花油茶的叶绿体基因组特征分析. 园艺学报, 47 (4):779-787.
[24]	Li Yongtan, Zhang Jun, Huang Yali, Fan Jianmin, Zhang Yiwen, Zuo Lihui. 2020. Analysis of chloroplast genome of Pyrus betulaefolia. Acta Horticulturae Sinica, 47 (6):1021-1032. (in Chinese)
	李泳潭, 张军, 黄亚丽, 范建敏, 张益文, 左力辉. 2020. 杜梨叶绿体基因组分析. 园艺学报, 47 (6):1021-1032.
[25]	Li Yu-nong. 1999. Research on the evolution of apple origin. Acta Horticulturae Sinica, 26 (4):213-220. (in Chinese)
	李育农. 1999. 苹果起源演化的考察研究. 园艺学报, 26 (4):213-220.
[26]	Lohse M, Drechsel O, Kahlau S, Bock R. 2013. Organellar Genome DRAW-a suite of tools for generating physical maps of plastid and mitochondrial genomes and visualizing expression data sets. Nucleic Acids Research, 41 (1):575-581. doi: 10.1093/nar/gks1075 URL
[27]	Lu M, Zhang H S, An H M. 2021. Chloroplast DNA-based genetic variation of Rosa roxburghii in Southwest China:phylogeography and conservation implications. Horticultural Plant Journal, 7 (4):286-294. doi: 10.1016/j.hpj.2021.01.002 URL
[28]	Ma B Q, Liao L, Peng Q, Fang T, Zhou H, Korban S S, Han Y P. 2017. Reduced representation genome sequencing reveals patterns of genetic diversity and selection in apple. Journal of Integrative Plant Biology, 59 (3):190-204. doi: 10.1111/jipb.12522
[29]	Ma B Q, Liao L, Zheng H Y, Chen J. 2015. Genes encoding aluminum-activated malate transporter II and their association with fruit acidity in apple. Plant Genome, 8:1-14.
[30]	Martin G, Cardi C. 2013. The complete chloroplast genome of banana(Musa acuminata,Zingiberales):insight into plastid monocotyledon evolution. PLoS ONE, 8 (6):e67350. doi: 10.1371/journal.pone.0067350 URL
[31]	Mccoy S R, Kuehl J V, Boore J L, Raubeson L A. 2008. The complete plastid genome sequence of Welwitschia mirabilis:an unusually compact plastome with accelerated divergence rates. BMC Evolutionary Biology, 8 (1):130-135. doi: 10.1186/1471-2148-8-130 URL
[32]	Nock C J, Waters D L, Edwards M A, Bowen S G, Rice N, Cordeiro G M, Henry R J. 2010. Chloroplast genome sequences from total DNA for plant identification. Plant Biotechnol, 9 (3):328-333. doi: 10.1111/j.1467-7652.2010.00558.x URL
[33]	Qian Jun. 2014. Study on chloroplast and mitochondrial genomes of Salvia miltiorrhiza[Ph. D. Dissertation]. Beijing:Peking Union Medical College. (in Chinese)
	钱俊. 2014. 丹参的叶绿体和线粒体基因组研究[博士论文]. 北京: 北京协和医学院.
[34]	Raubeson L A, Peery R, Chumley T W, Dziubek C, Fourcade H M, Boore J L, Jansen R K. 2007. Comparative chloroplast genomics:analyses including new sequences from the angiosperms Nuphar advena and Ranunculus macranthus. BMC Genomics, 8 (1):174-178. doi: 10.1186/1471-2164-8-174 URL
[35]	Ronquist F, Huelsenbeck J P. 2003. MrBayes3:Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12):1572-1574. doi: 10.1093/bioinformatics/btg180 pmid: 12912839
[36]	Sun X P, Jiao C, Schwaninger H, Thomas C, Ma Y M, Duan N B, Khan A, Fei Z J. 2020. Phased diploid genmoe assemblies and pan-genomes provide insights into the genetic history of apple domestication. Nature Genetics,1-10.
[37]	Velasco R, Zharikikh A, Affourtit J, Dhingra A, Cestaro A, Kalyanaraman A. 2010. The genome of the domesticated apple(Malus × domestica Borkh). Nature Genetics, 42 (10):833-839. doi: 10.1038/ng.654 URL
[38]	Wang R J, Cheng C L, Chang C C, Wu C L, Su T M, Chaw S M. 2008. Dynamics and evolution of the inverted repeat-large single copy junctions in the chloroplast genomes of monocots. BMC Evolutionary Biology, 36 (8):1471-2148.
[39]	Yan Guo-rong, Xu Zheng, Chu Wen-zhao. 2004. Study and protect on Malus sieversii. Beijing:Chinese Society for Horticultural Science:12-16. (in Chinese)
	阎国荣, 许正, 楚文照. 2004. 新疆野苹果(Malus sieversii)及其保护研究. 北京:中国园艺学会:12-16.
[40]	Yang Guo-feng, Su Kong-long, ZhaoYi-ran, Song Zhi-bin, Sun Juan. 2015. Analysis of codon usage in the chloroplast genome of Medicago truncatula. Acta Prataculturae Sinica, 24 (12):171-179. (in Chinese)
	杨国锋, 苏昆龙, 赵怡然, 宋智斌, 孙娟. 2015. 蒺藜苜蓿叶绿体密码子偏好性分析. 草业学报, 24 (12):171-179.
[41]	Zerbino D R, Birney E. 2008. Velvet:algorithms for de novo short read assembly using de Bruijn graphs. Genome Res, 18 (1):821-829. doi: 10.1101/gr.074492.107 URL
[42]	Zhang Li, Wang Yan, Chen Qing, Luo Ya,Zhang Yong,Tang Hao-ru,Wang Xiao-rong. 2015. Phylogenetic utility of Chinese Rubus(Rosaceae)based on ndhF sequence. Acta Horticulturae Sinica, 42 (1):19-30. (in Chinese)
	张丽, 王燕, 陈清, 罗娅, 张勇, 汤浩茹, 王小蓉. 2015. 基于ndhF序列的中国悬钩子属植物系统发育研究. 园艺学报, 42 (1):19-30.
[43]	Zheng Yi, Zhang Hui, Wang Qinmei, Gao Yue,Zhang Zhihong,Sun Yuxin. 2020. Complete chloroplast genome sequence of Clivia miniata and its characteristics. Acta Horticulturae Sinica, 47 (12):2439-2450. (in Chinese)
	郑祎, 张卉, 王钦美, 高悦, 张志宏, 孙玉新. 2020. 大花君子兰叶绿体基因组及其特征. 园艺学报, 47 (12):2439-2450.

基本特征 Basic characteristic	新疆野苹果 M. sieversii	‘元帅’ M.× domastica‘Red Delicious’	‘金冠’ M. × domastica‘Golden Delicious’
基因组Genomes/bp	160 230	160 288	160 068
LSC/bp	88 368	88 472	88 273
SSC/bp	19 196	19 174	19 181
IR/bp	26 333	26 321	26 370
蛋白编码基因Protein-coding genes	86	86	86
转运RNA基因tRNA gene	37	37	37
核糖体RNA基因rRNA gene	8	8	8
GC/%	36.5	36.5	36.6

基本特征 Basic characteristic	新疆野苹果 M. sieversii	‘元帅’ M.× domastica‘Red Delicious’	‘金冠’ M. × domastica‘Golden Delicious’
基因组Genomes/bp	160 230	160 288	160 068
LSC/bp	88 368	88 472	88 273
SSC/bp	19 196	19 174	19 181
IR/bp	26 333	26 321	26 370
蛋白编码基因Protein-coding genes	86	86	86
转运RNA基因tRNA gene	37	37	37
核糖体RNA基因rRNA gene	8	8	8
GC/%	36.5	36.5	36.6

区域 Regin	碱基 Base	新疆野苹果 M. sieversii	‘元帅’ M.× domastica‘Red Delicious’	‘金冠’ M. × domastica‘Golden Delicious’
大单拷贝区/% LSC	A	32.2	32.3	32.2
	C	17.6	17.6	17.6
	G	16.6	16.6	16.6
	T	33.6	33.6	33.6
	GC	34.1	34.1	34.2
小单拷贝区/% SSC	A	34.8	34.8	34.7
	C	16.0	16.0	15.9
	G	14.5	14.5	14.5
	T	34.8	34.8	34.8
	GC	30.4	30.4	30.4
反向重复区/% IRa	A	28.6	28.6	28.5
	C	22.1	22.1	22.1
	G	20.6	20.6	20.6
	T	28.7	28.8	28.8
	GC	42.7	42.7	42.7
总计/% Total	A	31.4	31.4	31.3
	C	18.6	18.6	18.6
	G	17.9	17.9	17.9
	T	32.1	32.1	32.1
	GC	36.5	36.5	36.6

区域 Regin	碱基 Base	新疆野苹果 M. sieversii	‘元帅’ M.× domastica‘Red Delicious’	‘金冠’ M. × domastica‘Golden Delicious’
大单拷贝区/% LSC	A	32.2	32.3	32.2
	C	17.6	17.6	17.6
	G	16.6	16.6	16.6
	T	33.6	33.6	33.6
	GC	34.1	34.1	34.2
小单拷贝区/% SSC	A	34.8	34.8	34.7
	C	16.0	16.0	15.9
	G	14.5	14.5	14.5
	T	34.8	34.8	34.8
	GC	30.4	30.4	30.4
反向重复区/% IRa	A	28.6	28.6	28.5
	C	22.1	22.1	22.1
	G	20.6	20.6	20.6
	T	28.7	28.8	28.8
	GC	42.7	42.7	42.7
总计/% Total	A	31.4	31.4	31.3
	C	18.6	18.6	18.6
	G	17.9	17.9	17.9
	T	32.1	32.1	32.1
	GC	36.5	36.5	36.6

基因组成 Group of genes	基因名称Gene name
基因组成 Group of genes	新疆野苹果M. sieversii	‘元帅’ M. × domastica‘Red Delicious’	‘金冠’ M. × domastica‘Golden Delicious’
核糖体RNA基因Ribosomal RNA genes	rrn16,rrn23,rrn4.5,rrn5	rrn23,rrn16,rrn4.5,rrn5	rrn23,rrn16,rrn4.5,rrn5
转运RNA基因 Transefer RNA genes	trnH-GUG,trnK-UUU,trnQ-UUG,trnS-GGU,trnG-UCC,trnR-UCU,trnC-GCA,trnD-GUC,trnY-GUA,trnT-GGU,trnS-UGA,trnG-GCC,trnfM-CAC,trnS-GGA,trnT-UGU,trnT-UAA,trnF-GAA,trnV-UAC,trnM-CAU,trnW-CCA,trnP-UGG,trnL-CAU,trnL-CAA,trnL-GAU,trnA-UGC,trnR-ACG,trnN-GUU,trnA-UGC,trnV-GAC,trnL-CAA	trnH-GUG,trnK-UUU,trnQ-UUG,trnS-GCU,trnG-UCC,trnR-UCU,trnC-GCA,trnD-GUC,trnE-UUC,trnT-GGU,trnS-UGA,trnfM-CAU,trnS-GGA,trnT-UGC,trnL-UAA,trnV-UAC,trnM-CAU,trnW-CCA,trnP-UGG,trnL-CAU,trnV-GAC,trnL-GAU,trnA-UGC,trnR-ACG,trnN-GUU,trnL-UAG,trnR-ACG,trnL-CAA,trnL-CAU	trnH-GUG,trnK-UUU,trnQ-UUG,trnS-GCU,trnG-UCC,trnR-UCU,trnC-GCA,trnD-GUC,trnY-GUA,trnE-UUC,trnT-GGU,trnS-UGA,trnfM-CAU,trnS-GGA,trnT-UGC,trnF-UGC,trnL-UAA,trnV-UAC,trnM-CAU,trnW-CCA,trnP-UGG,trnL-CAU,trnL-GAU,trnA-UGC,trnR-ACG,trnN-GUU,trnL-UAG,trnR-ACG,trnN-ACG,trnL-GAU,trnV-GAU,trnL-CAA,trnL-CAU
核糖体小亚基Small subunit of ribosom	rps16,rps2,rps14,rps4,rps8,rps7,rps19	rps16,rps19,rps2,rps14,rps4,rps12,rps11,rps8,rps7,rps15	rps16,rps14,rps4,rps18,rps8,rps12,rps3,rps7,rps15,rps19
核糖体大亚基 Large subunit of ribosom	rpl33,rpl20,rpl14,rpl16,rpl22,rpl12,rpl32,rpl23	rpL33,rpL20,rpL36,rpL14,rpL16,rpL22,rpL12,rpL23	rpL33,rpL20,rpL36,rpL14,rpL16,rpL23,rpL22,rpL38,rpL23,rpL2
DNA依赖型RNA聚合酶 DNA dependent RNA polymerase	rpoC1,rpoC2,rpoA	rpoC1,rpoC2,rpoB	rpoC1,rpoC2,rpoB
光合系统Ⅰ Subunit of photosestenⅠ	psa1,psa7,psaB	psaB,psaA,psaL,psaJ,psaC	psaA,psaB,psaL,psaJ,psaC
光合系统Ⅱ Subunit of photosestenⅡ	psbA,psbK,psbL,psbM,psbD,psbZ,psbJ,psbB,psbT,psbE,psbN	psbA,psbK,psbL,psbD,psbC,psbZ,psbJ,psbF,psbE,psbT,psbN,psbH,psbM	PsbB,psbK,psbL,psbM,psbD,psbC,psbZ,psbJ,psbL,psbF,psbE,psbT,psbH,psbN
细胞色素亚基Subunit of Cytochrome	petN,petA,petB,petD	petN,petA,petL,petG,petD,petB	petN,petA,petL,petG,petB,petD
ATP合成酶亚基Subunit of ATP synthase	atpE,atpA,atpH,atp1	atpA,atpF,atpL,atpE,atpB	atpA,atpF,atpH,atpL,atpB,atpE
ATP依赖型蛋白酶ATP dependent protease	clpP	clpP	clpP
Rubisco大亚基Large subunit of Rubisco	rbcL	rbcL	rbcL
NADH亚基 Subunit of NADH	ndhB,ndhF,ndhI,ndhJ,ndhC,ndhG,ndhE,ndhD,ndhH,ndhA,ndhK	ndhJ,ndhK,ndhC,ndhA,ndhG,ndhE,ndhD,ndhB,ndhF,ndhH,ndhI	ndhJ,ndhK,ndhC,ndhB,ndhF,ndhD,ndhA,ndhE,ndhG,ndhH,ndhI
成熟酶 Maturase	matK	matK	matK
外膜蛋白基因Envelop membrane protein	cemA	cemA	cemA
乙酰-CoA-羧基酶亚基 Subunit of Acetyl-CoA- Carboxyase	accD	accD	accD
C型细胞色素合成基因 C-type cytochrome synthesis gene	ccsA	ccsA	ccsA
保守的开放阅读框Conserved open reading frames	ycf3,ycf1,ycf4,ycf2	ycf3,ycf1,ycf4,ycf2	ycf3,ycf2,ycf1