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作物学报

作物学报 ›› 2021, Vol. 47 ›› Issue (6): 1124-1137.doi: 10.3724/SP.J.1006.2021.04150

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

花生油脂合成相关基因的表达谱分析

许静1(), 潘丽娟1, 李昊远2, 王通1, 陈娜1, 陈明娜1, 王冕1, 禹山林1, 侯艳华2,*(), 迟晓元1,*()   

  1. 1山东省花生研究所, 山东青岛 266100
    2哈尔滨工业大学(威海)海洋科学与技术学院, 山东威海 264209
  • 收稿日期:2020-07-10 接受日期:2020-11-13 出版日期:2021-06-12 网络出版日期:2020-12-23
  • 通讯作者: 侯艳华,迟晓元
  • 作者简介:许静, E-mail:xu_jing_cool@yeah.net
  • 基金资助:
    国家现代农业产业技术体系(花生)建设专项(CARS-13);青岛市科技惠民示范引导专项(20-3-4-26-nsh);山东省农业科学院农业科技创新工程(CXGC2016B02);山东省农业科学院农业科技创新工程(CXGC2018E21);山东省重大科技创新工程项目(2019JZZY010702);广东省重点领域研发计划项目(2020B020219003);山东省良种工程(2017LZGC003);临沂市重点研发计划项目(2019YD009);国家自然科学基金项目(31701464)

Expression pattern analysis of genes related to lipid synthesis in peanut

XU Jing1(), PAN Li-Juan1, LI Hao-Yuan2, WANG Tong1, CHEN Na1, CHEN Ming-Na1, WANG Mian1, YU Shan-Lin1, HOU Yan-Hua2,*(), CHI Xiao-Yuan1,*()   

  1. 1Shandong Peanut Research Institute, Qingdao 266100, Shandong, China
    2School of Marine and Technology, Harbin Institute of Technology, Weihai 264209, Shandong, China
  • Received:2020-07-10 Accepted:2020-11-13 Published:2021-06-12 Published online:2020-12-23
  • Contact: HOU Yan-Hua,CHI Xiao-Yuan
  • Supported by:
    The China Agriculture Research System(CARS-13);The Qingdao People’s livelihood Science and Technology Program(20-3-4-26-nsh);The Agricultural Scientific and Technological Innovation Project of Shandong Academy of Agricultural Sciences(CXGC2016B02);The Agricultural Scientific and Technological Innovation Project of Shandong Academy of Agricultural Sciences(CXGC2018E21);The Breeding Project from Department Science & Technology of Shandong Province(2019JZZY010702);The Key-Area Research and Development Program of Guangdong Province(2020B020219003);The Improved Variety Engineering Project of Shandong Province(2017LZGC003);The Key Research and Development Program of Linyi(2019YD009);The National Natural Science Foundation of China(31701464)

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摘要：

为研究不同发育时期花生籽仁油脂合成过程中基因表达调控模式, 本研究以高油酸、中油花生品系F18和低油酸、低油花生品种‘鲁花6号’为材料, 对下针后10、30、40、60 DAP (days after pegging)的花生种子进行表达谱芯片测序。结果表明, 130、3556、2783个基因分别在30、40、60 DAP时期差异表达。GO注释和KEGG富集结果显示, 差异表达的基因主要富集在脂肪酸合成和光合等代谢进程中, 其中FAB2、FAD2、WRI1等主要参与油酸的积累, 参与光合作用的基因均为捕光叶绿素a/b结合蛋白, 全部上调表达。代谢通路图结果表明, 籽仁发育的40 DAP和60 DAP时期, 脂肪酸合成途径的基因均上调表达。研究结果为花生油脂代谢的分子机制提供理论基础, 同时也为花生品质改良贡献了基因资源。

关键词: 花生, 基因芯片, 差异表达分析, 油脂合成相关基因

Abstract:

In order to survey the regulation patterns of genes expression in the synthesis of oil at different developmental stages in peanut seed, F18 (high-oleic medium oil peanut variety) and ‘Luhua 6’ (low-oleic acid low oil variety) were used as research materials. The expression pattern analysis of genes was performed for peanut seeds on the 10, 30, 40, and 60 DAP, respectively. The results indicated that 130, 3556, and 2783 genes were significantly differentially expressed in the two varieties (lines) on the 30, 40, and 60 DAP, respectively. GO annotation and KEGG enrichment showed that DEGs were mainly enriched in the fatty acid synthesis and photosynthetic pathways, such as FAB2, FAD2, WRI1 genes, which were involved in the accumulation of oleic acid. All the genes involved in photosynthesis pathway were photochlorophyll binding proteins and all of them were up-regulated. KEGG pathway indicated that all the genes involved in fatty acid biosynthesis pathway on the 40 DAP and 60 DAP were up-regulated. In summary, these results provide a theoretical basis for molecular study of fatty acid synthesis in the development stage of peanut seeds and offer some candidate genes as gene resources of quality improvement in peanut breeding.

Key words: peanut, microarray, differentially expressed genes, genes involved in lipid synthesis

表1

RT-PCR验证基因所用的引物"

基因编号
Gene ID		 正向引物
Forward primer (5°-3°)		 反向引物
Reverse primer (5°-3°)
AHTC1032413 ACGAAGGTTATGGAAGTAG AGGAAGGTAGTGAAGGAA
AHTC1030930 AGGTGAAGAATCTGTTAGTC CTTATCTTGTGAATTACTTGTCT
AHTC1005724 GATAAGTGGAGCAGTAGC CAAGAAGACGATGATATAAGC
AHTC1030772 TCAATCAGATTCGTAACATTC ACTCAATAACTCTATCCACTAG
AHTC1020107 CTACTACTCATCATTCATTGTC CTTAACTCACCCACACAT
AHTC1012845 TCCTCTTCTTCATCTTCATC CCTGGTTACTCCTCTGTA
AHTC1014100 ACTCCTTCTCCTCTTCAT ATATTAGGCAGCAGATACC
AHTC1001812 AATGGCAGGAGGTTATTG TTCTTCTTCATCATAATCTTAGC
AHTC1001844 GTGACTTGGCAGAGATTA AATTGATTGATTGACGGATG
AHTC1001749 TCTTGGAGTTCCTTGTTG GCTATGCGACTTCTTCAT
AHTC1005424 GAGTGATGTAGTTGAGATAGA CTCTTCCGATGACTTGAT
AHTC1000715 TGGTTAAGAGGAGCATTG ATGAGGCATTGTTGAGAA
AHTC1002482 GTTGCTCTGATGACTCTC TATGGCTAACCGTTGATG
AHTC1027178 TCTCACTCACTCTCATCA CCACCACCATCTTAGAAG
Actin11 TTGGAATGGGTCAGAAGGATGC AGTGGTGCCTCAGTAAGAAGC

表2

不同发育时期两品种(系)油脂和脂肪酸含量"

样品名称
Name		 发育时期
Development stage		 含油量
Oil content (%)		 油酸含量
Oleic content (%)		 亚油酸含量
Linoleic content (%)		 油亚比
O/L
F18 10 DAP 13.86 58.91 15.39 3.83
30 DAP 35.21 74.01 5.61 13.19
40 DAP 46.82 76.06 2.43 31.30
60 DAP 50.81 70.75 1.55 45.65
鲁花6号
Luhua 6		 10 DAP 19.67 31.63 37.14 0.85
30 DAP 36.95 34.96 41.28 0.85
40 DAP 44.44 37.67 39.39 0.96
60 DAP 45.44 43.98 34.28 1.28

图1

不同发育时期两品种(系)间差异表达基因火山图 A: 10 DAP差异表达基因; B: 30 DAP差异表达基因; C: 40 DAP差异表达基因; D: 60 DAP差异表达基因。红色表示F18相对于‘鲁花6号’差异表达的基因。"

图2

不同品种(系)间不同发育时期共同表达的差异基因 A: F18相比于‘鲁花6号’上调表达基因; B: F18相比于‘鲁花6号’下调表达基因。"

图3

差异表达基因GO注释及富集分析"

图4

差异表达基因KEGG注释及富集分析"

图5

40 DAP和60 DAP差异表达基因脂肪酸代谢通路图 红色代表相对于‘鲁花6号’, F18上调表达的基因。"

表3

40 DAP发育时期油脂代谢相关基因"

基因编号
Gene ID		 基因注释
Gene annotation		 GO富集
GO enrichment		 调控a
Regulationa		 差异表达倍数
Fold change
AHTC1031703 移酶1-酰基-sn-甘油-3-磷酸酰基转 磷脂生物合成 上调 4.19
1-acyl-sn-glycerol-3-phosphate acyltransferase-like Phospholipid biosynthetic process Up
基因编号
Gene ID		 基因注释
Gene annotation		 GO富集
GO enrichment		 调控a
Regulationa		 差异表达倍数
Fold change
AHTC1022314 3-酮脂酰-合酶 脂肪酸生物合成 上调 2.88
3-ketoacyl-synthase Fatty acid biosynthetic process Up
AHTC1029564 类3-酮脂酰-合酶10 脂肪酸生物合成 上调 2.30
3-ketoacyl-synthase 10-like Fatty acid biosynthetic process Up
AHTC1007527 3-酮脂酰-合酶11 脂肪酸生物合成 下调 3.28
3-ketoacyl-synthase 11 Fatty acid biosynthetic process Down
AHTC1008395 类3-酮脂酰-合酶11 脂肪酸生物合成 下调 2.55
3-ketoacyl-synthase 11-like Fatty acid biosynthetic process Down
AHTC1016489 3-酮脂酰-合酶21 脂质生物合成 上调 5.25
3-ketoacyl-synthase 21 Lipid biosynthetic process Up
AHTC1003640 类3-酮脂酰-合酶21 脂肪酸生物合成 上调 3.51
3-ketoacyl-synthase 21-like Fatty acid biosynthetic process Up
AHTC1003641 类3-酮脂酰-合酶21 脂肪酸生物合成 上调 6.43
3-ketoacyl-synthase 21-like Fatty acid biosynthetic process Up
AHTC1004544 3-酮脂酰-合酶6 脂肪酸生物合成 上调 2.86
3-ketoacyl-synthase 6 Fatty acid biosynthetic process Up
AHTC1019082 α-乙酰羧化酶亚基 脂肪酸生物合成; 丙二酰辅酶A生物合成 下调 2.07
Alpha-acetyl-carboxylase subunit Fatty acid biosynthetic process;
malonyl-CoA biosynthetic process		 Down
AHTC1001812 β-酮酰基-ACP还原酶 脂肪酸生物合成; 氧化还原 上调 2.05
Beta-ketoacyl-acp reductase 1-2 Fatty acid biosynthetic process;
oxidation-reduction process		 Up
AHTC1001844 β-酮酰基-ACP合酶 脂肪酸生物合成 上调 2.17
Beta-ketoacyl-acp synthase i-2 Fatty acid biosynthetic process Up
AHTC1005915 叶绿体ω-6 脂肪酸去饱和酶 二磷酸生物合成 上调 2.06
Chloroplast omega-6 fatty acid desaturase Diphosphate biosynthetic process Up
AHTC1032453 酰基-载体-蛋白合酶 脂肪酸生物合成 上调 2.16
Fatty acid biosynthesis 1 isoform 1 Fatty acid biosynthetic process Up
AHTC1032454 酰基-载体-蛋白合酶 脂肪酸生物合成 上调 2.57
Fatty acid biosynthesis 1 isoform 1 Fatty acid biosynthetic process Up
AHTC1020107 鞘脂Δ8 去饱和酶 脂质代谢; 氧化还原 上调 2.04
Fatty acid sphingolipid desaturase Lipid metabolic process; oxidation-
reduction process		 Up
AHTC1035094 脂质转移蛋白 脂质转移 上调 14.59
Lipid transfer protein gpi-anchored Lipid transport Up
AHTC1009675 长链酰基合成酶 长链脂肪酸代谢 上调 2.44
Long chain acyl-synthetase 1-like isoform×1 Long-chain fatty acid metabolic process Up
AHTC1026013 长链酰基合成酶 长链脂肪酸代谢 上调 3.03
Long chain acyl-synthetase 1-like isoform×1 Long-chain fatty acid metabolic process up
AHTC1017689 长链酰基合酶2 长链脂肪酸代谢 上调 2.65
Long chain acyl-synthetase 2 Long-chain fatty acid metabolic process up
AHTC1006429 长链酰基合成酶2异构体 长链脂肪酸代谢 上调 3.44
Long chain acyl-synthetase 2-like isoform×2 Long-chain fatty acid metabolic process Up
AHTC1034617 微粒体ω-3脂肪酸去饱和酶 脂质代谢; 氧化还原 上调 5.40
Microsomal omega-3 fatty acid desaturase Lipid metabolic process; oxidation-
reduction process		 Up
基因编号
Gene ID		 基因注释
Gene annotation		 GO富集
GO enrichment		 调控a
Regulationa		 差异表达倍数
Fold change
AHTC1020991 非特异性脂质转移酶蛋白 脂质转移 上调 3.92
Non-specific lipid-transfer protein Lipid transport Up
AHTC1017861 ω-3去饱和酶家族蛋白 不饱和脂肪酸生物合成; 氧化还原 下调 2.55
Omega-3 desaturase family protein Unsaturated fatty acid biosynthetic process; oxidation-reduction process Down
AHTC1008662 硬脂酰ACP去饱和酶 脂肪酸生物合成 上调 3.48
Stearoyl-acyl-carrier desaturase Fatty acid biosynthetic process; oxidation-reduction process Up

表4

60 DAP发育时期油脂代谢相关基因"

基因编号
Gene ID		 基因注释
Gene annotation		 GO富集
GO enrichment		 调控a
Regulationa		 差异表达倍数
Fold change
AHTC1005431 类3-酮脂酰-合酶11 脂肪酸生物合成 上调 7.27
3-ketoacyl-synthase 11-like Fatty acid biosynthetic process Up
AHTC1000075 酰基载体蛋白 脂肪酸生物合成; 氧化还原 上调 3.10
Acyl carrier protein 1 Fatty acid biosynthetic process; oxidation- reduction process Up
AHTC1000128 酰基载体蛋白 脂肪酸生物合成; 氧化还原 上调 2.72
Acyl carrier protein 1 Fatty acid biosynthetic process; oxidation- reduction process Up
AHTC1028756 α-羧基转移酶前体 脂肪酸生物合成 上调 4.68
Alpha-carboxyltransferase precursor Fatty acid biosynthetic process Up
AHTC1005278 α-羧基转移酶 脂肪酸生物合成; 泛醌生物合成 上调 4.26
Alpha-carboxyl transferase 2 Fatty acid biosynthetic process; coenzyme biosynthetic process Up
AHTC1000881 β-羟烷基-ACP脱氢酶 脂肪酸生物合成 上调 18.60
Beta-hydroxyacyl-acp dehydratase Fatty acid biosynthetic process Up
AHTC1001812 β-酮酰基-ACP还原酶 脂肪酸生物合成; 氧化还原 上调 7.89
Beta-ketoacyl-acp reductase 1-2 Fatty acid biosynthetic process; oxidation- reduction process Up
AHTC1001844 β-酮酰基-ACP合酶 脂肪酸生物合成 上调 4.49
Beta-ketoacyl-acp synthase i-2 Fatty acid biosynthetic process Up
AHTC1007010 β-酮酰基-ACP合酶 脂肪酸生物合成 上调 3.25
Beta-ketoacyl-acp synthase ii-1 Fatty acid biosynthetic process Up
AHTC1002452 生物素羧基载体蛋白 脂肪酸生物合成 上调 6.29
Biotin carboxyl carrier protein 2-2 Fatty acid biosynthetic process Up
AHTC1014935 生物素羧基载体蛋白 脂肪酸生物合成 上调 2.83
Biotin carboxyl carrier protein 2-2 Fatty acid biosynthetic process Up
AHTC1022457 生物素羧基载体蛋白 脂肪酸生物合成 上调 4.01
Biotin carboxyl carrier protein 2-2 Fatty acid biosynthetic process Up
AHTC1032453 酰基-载体-蛋白合酶 脂肪酸生物合成 上调 3.90
Fatty acid biosynthesis 1 isoform 1 Fatty acid biosynthetic process Up
AHTC1009675 长链酰基合成酶 长链脂肪酸代谢 上调 7.57
Long chain acyl-synthetase 1-like isoform×1 Long-chain fatty acid metabolic process Up
基因编号
Gene ID		 基因注释
Gene annotation		 GO富集
GO enrichment		 调控a
Regulationa		 差异表达倍数
Fold change
AHTC1026013 长链酰基合成酶 长链脂肪酸代谢 上调 9.64
Long chain acyl-synthetase 1-like isoform×1 Long-chain fatty acid metabolic process Up
AHTC1017689 长链酰基合酶2 长链脂肪酸代谢 上调 6.75
Long chain acyl-synthetase 2 Long-chain fatty acid metabolic process Up
AHTC1020991 非特异性脂质转移酶蛋白 脂质转移 上调 3.89
Non-specific lipid-transfer protein Lipid transport Up
AHTC1002315 非特异性脂质转移酶蛋白 脂质转移 上调 3.73
Non-specific lipid-transfer protein at5g64080-like Lipid transport Up
AHTC1015374 非特异性脂质转移酶蛋白 脂质转移 上调 3.13
Non-specific lipid-transfer protein at5g64080-like Lipid transport Up
AHTC1019533 酰基转移酶 甘油酯生物合成 下调 6.23
O-acyltransferase wsd1-like Glycerolipid biosynthetic process Down
AHTC1005424 油酰-酰基载体蛋白硫酯酶 脂肪酸生物合成 上调 2.72
Oleoyl-acyl carrier protein thioesterase chloroplastic-like Fatty acid biosynthetic process Up
AHTC1000715 ω6 脂肪酸去饱和酶 脂质代谢; 氧化还原 上调 3.82
Omega 6 fatty acid desaturase Lipid metabolic process; oxidation-reduction process Up
AHTC1021791 过氧化物酶体脂肪酸-β氧化多功能蛋白 脂肪酸生物合成; 氧化还原 下调 2.62
Peroxisomal fatty acid beta-oxidation multifunctional protein mfp2-like Fatty acid metabolic process; oxidation-
reduction process		 Down
AHTC1001749 硬脂酰-ACP去饱和酶 脂肪酸生物合成; 氧化还原 上调 29.89
Searoyl-acp desaturase Fatty acid biosynthetic process; oxidation- reduction process Up
AHTC1003174 超长链3-酰基还原酶 超长链脂肪酸生物合成 上调 3.92
Very-long-chain 3-oxoacyl-reductase 1-like Very long-chain fatty acid biosynthetic process Up
AHTC1001068 类超长链烯酰还原酶异构体 脂质代谢; 氧化还原 上调 3.97
Very-long-chain enoyl-reductase-like isoformx1 Lipid metabolic process; oxidation-reduction process Up
AHTC1001069 类超长链烯酰还原酶异构体 脂质代谢; 氧化还原 上调 3.80
Very-long-chain enoyl-reductase-like isoformx1 Lipid metabolic process; oxidation-reduction process Up

图6

WRI1蛋白互作网络 黑色和灰色分别代表相对于‘鲁花6号’, F18上调和下调表达的基因。"

图7

不同发育时期差异表达基因的RT-PCR验证结果"

表5

差异表达基因的RT-PCR验证结果"

基因编号
Gnen ID		 基因注释
Gene annotation		 发育时期
Development stage		 调控a
Regulation a		 芯片数据(倍数)
Microarray data
(fold change)		 RT-PCR验证结果(倍数)
RT-PCR result
(fold change)
AHTC1032413 黄酮醇合酶 DAP30 上调 6.05 13.64
Flavonol synthase flavanone 3-hydroxylase-like Up
AHTC1030930 锌指蛋白 DAP30 下调 -3.78 -4.55
Dof zinc finger protein Down
AHTC1005724 非特异性脂质转移酶 DAP40 上调 4.04 3.25
Non-specific lipid-transfer protein 3-like Up
AHTC1030772 β-羟酰基-ACP脱水酶 DAP40 上调 2.86 2.11
Beta-hydroxyacyl-acp dehydrase 1 Up
AHTC1020107 鞘脂Δ8 去饱和酶 DAP40 上调 2.04 3.48
Fatty acid sphingolipid desaturase Up
AHTC1012845 WRI1 转录因子 DAP40 上调 2.15 2.31
Ethylene-responsive transcription factor wri1-like Up
AHTC1030930 锌指蛋白 DAP40 下调 -5.30 -10.24
Dof zinc finger protein down
AHTC1014100 肌醇半乳糖醇合酶 DAP40 下调 -3.09 -4.07
Galactinol synthase 8 Down
AHTC1001812 β-酮酰基-ACP还原酶 DAP60 上调 7.89 5.09
Beta-ketoacyl-acp reductase 1-2 Up
AHTC1001844 β-酮酰基-ACP合酶 DAP60 上调 4.49 4.83
Beta-ketoacyl-acp synthase i-2 Up
AHTC1012845 WRI1 转录因子 DAP60 上调 5.83 21.21
Ethylene-responsive transcription factor wri1-like Up
AHTC1005424 酰基载体蛋白硫酯酶 DAP60 上调 2.72 3.47
Oleoyl-acyl carrier protein thioesterase chloroplastic-like Up
基因编号
Gnen ID		 基因注释
Gene annotation		 发育时期
Development stage		 调控a
Regulationa		 芯片数据(倍数)
Microarray data
(fold change)		 RT-PCR验证结果(倍数)
RT-PCR result
(fold change)
AHTC1001749 硬脂酰-ACP去饱和酶 DAP60 上调 29.89 16.26
Stearoyl-acp desaturase Up
AHTC1000715 ω6 脂肪酸去饱和酶 DAP60 上调 3.82 1.55
Omega 6 fatty acid desaturase Up
AHTC1002482 天冬酰胺合成酶 DAP60 下调 -4.87 -6.12
Asparagine synthetase 2 Down
AHTC1027178 Hsf 转录因子 DAP60 下调 -4.39 -9.85
Heat stress transcription factor a-2-like Down
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