中国农业科学 ›› 2019, Vol. 52 ›› Issue (9): 1587-1600.doi: 10.3864/j.issn.0578-1752.2019.09.010
龚成胜,赵胜杰,路绪强,何楠,朱红菊,豆峻岭,袁平丽,李兵兵,刘文革()
收稿日期:
2018-10-28
接受日期:
2018-02-17
出版日期:
2019-05-01
发布日期:
2019-05-16
通讯作者:
刘文革
作者简介:
龚成胜,E-mail: 基金资助:
GONG ChengSheng,ZHAO ShengJie,LU XuQiang,HE Nan,ZHU HongJu,DOU JunLing,YUAN PingLi,LI BingBing,LIU WenGe()
Received:
2018-10-28
Accepted:
2018-02-17
Online:
2019-05-01
Published:
2019-05-16
Contact:
WenGe LIU
摘要:
【目的】蜡粉是植物抵御外界胁迫的第一层保护性屏障,对西瓜果实表皮蜡粉的结构、化学成分、遗传规律进行研究,并预测控制该性状的基因,以便全面了解性状的生理生化作用,发掘候选基因。【方法】试验选用无蜡粉的西瓜自交系‘美佳选黑’(P1)和有蜡粉的西瓜自交系‘FH’(P2)为双亲配制杂交组合,构建六世代群体(P1、P2、F1、F2、BC1P1、BC1P2),利用扫描电子显微镜(SEM)对双亲成熟期果实表皮结构进行观察,通过气相色谱-质谱联用仪(GC-MS)对蜡粉的化学成分进行测定,以峰面积为指标,定量计算蜡粉化学成分的含量,采用BSA-seq对西瓜表皮蜡粉进行基因初定位,应用BLAST软件将定位区间内的编码基因与多个数据库比对完成基因信息注释,并通过详细的基因注释信息及突变位点分析,快速筛选候选基因。【结果】西瓜果实表皮蜡粉为灰白色,呈致密的板状结构,长度约为5 μm;无蜡粉材料表皮较为光滑,无蜡粉层附着。GC-MS分析显示,蜡粉中共检测到24种脂肪族化学成分,分别属于烃类、醇类、酯类、酸类、酚类和醛类。包括烃类物质10种,含量占表皮蜡粉有效化学提取物总含量的77.72%,链长变化范围为C17—36,主要为C27、C28、C29、C32、C33、C34、C36的饱和正链烷烃;醇类物质5种,占12.60%;酯类物质5种,占0.43%;酸类物质2种,占0.53%;酚类1种,占0.80%;醛类1种,占3.99%;含量最高的5种化学成分依次为:正三十四烷、正二十九烷、1,30-三十烷二醇、正三十三烷、正二十八烷。在后代群体中,F1、BC1P2群体西瓜果实表皮全部有蜡粉,F2群体有蜡粉和无蜡粉西瓜果实的分离比符合3﹕1的孟德尔分离比例,BC1P1回交群体有蜡粉和无蜡粉分离比符合1﹕1的理论比,说明蜡粉的有无符合单基因显性遗传模式,有蜡粉对无蜡粉为显性。对BSA-seq数据进行SNP和InDel关联分析,关联结果取交集得到1号染色体3.16—4.84 Mb的候选区域,该区域共含有144个基因。数据库比对结果显示,在关联区域中,共138个基因有功能注释,包括10个非同义突变,1个移码突变。结合已有文献报道,其中5个非同义突变基因可能与西瓜表皮蜡粉的生成有关:Cla002367为烯酰ACP-还原酶(ECR)类基因,该类基因是特长链脂肪酸合成所必须;Cla011514、Cla002337和Cla002342为细胞色素P450(CYP)家族基因,该家族部分编码蛋白能够通过烃基羟化等反应催化脂肪族化合物的生成;Cla002353的基因注释信息为ABC转运体,ABC转运体与蜡质分子的转运息息相关。【结论】西瓜果实表皮蜡粉为板状结构,主要由特长链脂肪酸衍生成的脂肪族化合物组成。该性状是单基因遗传,有蜡粉为显性性状。BSA关联分析得到1号染色体1.68 Mb的关联区域,并预测关联区域中Cla002367、Cla011514、Cla002337、Cla002342、Cla002353这5个非同义突变基因为调控西瓜果实表皮蜡粉性状的候选基因。
龚成胜, 赵胜杰, 路绪强, 何楠, 朱红菊, 豆峻岭, 袁平丽, 李兵兵, 刘文革. 西瓜果实表皮蜡粉的化学成分与基因定位[J]. 中国农业科学, 2019, 52(9): 1587-1600.
GONG ChengSheng, ZHAO ShengJie, LU XuQiang, HE Nan, ZHU HongJu, DOU JunLing, YUAN PingLi, LI BingBing, LIU WenGe. Chemical Compositions and Gene Mapping of Wax Powder on Watermelon Fruit Epidermis[J]. Scientia Agricultura Sinica, 2019, 52(9): 1587-1600.
表1
西瓜果实表皮蜡粉的组成成分与相对含量"
化合物类别 Categories | 化合物名称 Compounds | 保留时间 Retention time (min) | 相对含量 Relative content (%) |
---|---|---|---|
正二十四烷* Tetracosane* | 17.20 | ||
烃类 Hydrocarbons | 2,6,10-三甲基十四烷 Tetradecane,2,6,10-trimethyl- | 4.92 | 0.05 |
3-乙基-5-(2-乙基丁基)十八烷 Octadecane, 3-ethyl-5-(2-ethylbutyl)- | 19.00 | 0.01 | |
正二十八烷 Octacosane | 23.45 | 5.89 | |
正二十九烷 Nonacosane | 27.92 | 16.24 | |
17-三十五烯 17-Pentatriacontene | 28.58 | 0.60 | |
正二十七烷 Heptacosane | 29.90 | 1.14 | |
正三十四烷 Tetratriacontane | 32.31 | 40.76 | |
正三十二烷 Dotriacontane | 34.02 | 1.07 | |
正三十三烷 Tritriacontane | 36.12 | 10.67 | |
正三十六烷 Hexatriacontane | 39.79 | 0.93 | |
酯类 Esters | (E)-3,7-二甲基-2,6-辛二烯醇3-甲基丁酸酯 Geranyl isovalerate | 5.28 | 0.10 |
顺式-9-十八碳烯酸-2-苯基-1,3-二氧杂环己烷-4-甲基酯 9-Octadecenoic acid, (2-phenyl-1,3-dioxolan-4-yl)methyl ester, cis- | 7.58 | 0.09 | |
俞酸甘油酯 Docosanoic acid, 1,2,3-propanetriyl ester | 35.83 | 0.11 | |
胆酸乙酯 Ethyl iso-allocholate | 12.21 | 0.05 | |
十八烯酸-3(十八烷氧基)丙酯 Oleic acid, 3-(octadecyloxy)propyl ester | 40.79 | 0.07 | |
醇类 Alcohols | 1-三十七烷醇 1-Heptatriacotanol | 9.64 | 0.15 |
E,E,Z-1,3,12-十九烯-4,14-二醇 E,E,Z-1,3,12-Nonadecatriene-4,14-diol | 13.04 | 0.21 | |
1,30-三十烷二醇 1,30-Triacontanediol | 34.99 | 11.55 | |
1-三十五醇 1-Pentatriacontanol | 36.86 | 0.48 | |
乙酰氧基-7,8-环氧羊毛甾烷-11-醇 7,8-Epoxylanostan-11-ol, 3-acetoxy- | 37.58 | 0.21 | |
酸类 Acids | 顺式-1-二十碳烯酸 cis-13-Eicosenoic acid | 6.83 | 0.33 |
五氟丙酸八碳五烯酸Octatriacontyl pentafluoropropionate | 40.29 | 0.20 | |
酚类Phenols | 2,4-二叔丁基苯酚Phenol,2,4-bis(1,1-dimethylethy)- | 5.701 | 0.80 |
醛类Aldehyde | 十八醛 Octadecanal | 26.42 | 3.99 |
图6
候选区域内基因的KEGG功能分类 C1:内吞作用;C2:自噬的调节;G1:基底切除修复;G2:DNA复制;G3:同源重组;G4:错配修复;G5:核苷酸切除修复;G6:氨酰-tRNA生物合成;G7:核糖体;M1:半胱氨酸和蛋氨酸代谢;M2:甘氨酸、丝氨酸、苏氨酸代谢;M3:苯丙氨酸代谢;M4:酪氨酸代谢;M5:异喹啉生物碱生物合成;M6:对苯二甲酸、哌啶和吡啶生物碱的生物合成;M7:氨基糖和核苷酸糖代谢;M8:果糖和甘露糖代谢;M9:半乳糖代谢;M10:糖酵解/糖异生;M11:戊糖磷酸途径;M12:丙酸盐代谢;M13:丙酮酸代谢;M14:淀粉和蔗糖代谢;M15:氧化磷酸化;M16:氨基酸的生物合成;M17:碳代谢;M18:脂肪酸代谢;M19:脂肪酸生物合成;M20:甘油磷酸代谢;M21:β-丙氨酸代谢;M22:二萜生物合成;M23:嘌呤代谢;M24:嘧啶代谢"
表3
候选区域内基因功能注释结果统计"
功能注释数据库Annotated_databases | 有注释信息基因数 Annotated information gene number | 存在非同义突变的基因数Number of genes with nonsynonymous mutations | 存在移码突变的基因数 Number of genes with frameshift mutation |
---|---|---|---|
GO | 42 | 6 | 0 |
KEGG | 46 | 4 | 0 |
COG | 63 | 8 | 1 |
NR | 137 | 10 | 1 |
NT | 133 | 10 | 1 |
trEMBL | 136 | 10 | 1 |
SwissProt | 118 | 10 | 1 |
Total | 138 | 10 | 1 |
表4
候选区域内非同义突变基因和移码突变基因功能注释"
突变类型 Mutation type | 基因 Gene Id | 基因注释 Gene annotation |
---|---|---|
非同义突变 Non synonymous mutation | Cla011514 | 细胞色素P450 Cytochrome P450 |
Cla011577 | 乳酸/苹果酸脱氢酶 Lactate/malate dehydrogenase, | |
Cla011542 | 铜胺氧化酶,酶结构域 Copper amine oxidase, enzyme domain | |
Cla011561 | 3-Oxoacyl-[酰基载体蛋白(ACP)]合成酶Ⅲ 3-Oxoacyl-[acyl-carrier-protein (ACP)] synthase III | |
Cla002342 | 含IPRO结构域(S)的IPRO1128细胞色素P450 Contains Inerpro domain(s) IPR001128 Cytochrome P450 | |
Cla002367 | 烯酰基-(酰基载体蛋白)还原酶 Enoyl-(Acyl carrier protein) reductase | |
Cla002326 | 木聚糖酶抑制剂N端 Xylanase inhibitor N-terminal | |
Cla002353 | ABC转运体 ABC transporter | |
Cla002337 | 细胞色素P450 Cytochrome P450 | |
Cla011554 | 无顶端分生组织(NAM)蛋白 No apical meristem (NAM) protein | |
移码突变 Frameshift mutation | Cla002327 | rRNA/tRNA 2′-O-甲基转移酶类纤维蛋白样蛋白 rRNA/tRNA 2'-O-methyltransferase fibrillarin-like protein |
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