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作物学报 ›› 2019, Vol. 45 ›› Issue (3): 381-389.doi: 10.3724/SP.J.1006.2019.84105

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蓝型油菜皖油20号种子不同部位油脂合成的转录调控分析

张宇婷,鲁少平,金诚,郭亮()   

  1. 华中农业大学作物遗传改良国家重点实验室, 湖北武汉 430070
  • 收稿日期:2018-07-28 接受日期:2018-12-24 出版日期:2019-03-12 网络出版日期:2019-01-03
  • 通讯作者: 郭亮
  • 基金资助:
    本研究由国家自然科学基金青年科学基金项目(31701458);中央高校基本科研业务费专项资金资助项目资助(2662015PY090)

Transcriptional regulation of oil biosynthesis in different parts of Wanyou 20 (Brassica napus) seeds

Yu-Ting ZHANG,Shao-Ping LU,Cheng JIN,Liang GUO()   

  1. National Key Laboratory of Crop Genetic Improvement, Huazhong Agricultural University, Wuhan 430070, Hubei, China
  • Received:2018-07-28 Accepted:2018-12-24 Published:2019-03-12 Published online:2019-01-03
  • Contact: Liang GUO
  • Supported by:
    This study was supported by the National Science Foundation for Young Scientists of China(31701458);Fundamental Research Funds for the Central Universities(2662015PY090)

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摘要：

甘蓝型油菜是主要的油料作物之一, 种子含油量一般在35%~50%。油脂主要储存于油菜种子胚中, 胚主要由子叶[包括外子叶(OC)和内子叶(IC)和胚轴(EA)]组成。低芥酸油菜品种皖油20号(WY20)种子不同部位的含油量存在显著差异。WY20的胚中, OC含油量最高, EA含油量最低。同时, 脂肪酸组成在种子不同部位也存在差异, EA中棕榈酸(C16:0)、亚油酸(C18:2)及二十碳酸(C20:0)的比例均显著高于子叶, 特别是C16:0在EA中的比例约为子叶的2倍。而油酸(C18:1)及二十碳烯酸(C20:1)在子叶中的比例均显著高于EA。硬脂酸(C18:0)在OC中含量最低, 在IC和EA中无差别。亚麻酸(C18:3)则在OC中含量最高, 在IC和EA中无差异。对发育34d种子的IC、OC和EA进行转录组分析, 将三个部位中基因表达定量分析的结果两两比较后共发掘出7192个差异表达基因, 其中OC和IC之间差异表达基因数目较少, 子叶和EA间有较多的差异表达基因。子叶和胚轴中的差异表达基因富集在光合作用、脂肪酸代谢和叶绿素合成等生物学过程。基因功能注释显示, 差异表达基因中有355个和脂质代谢相关, 且多集中在质体中脂肪酸从头合成途径。本研究表明油脂合成途径关键基因的差异调控是造成油菜种子不同部位含油量和脂肪酸组成差异的主要因素。

关键词: 甘蓝型油菜, 种子不同部位, 含油量, 脂肪酸组成, 转录调控

Abstract:

Brassica napus is one of the main oil crops and the seed oil content is generally between 35% and 50%. Oil is mainly stored in the seed embryo. Embryo is composed of cotyledons (including outer and inner cotyledons) and embryonic axis. The oil content and fatty acid composition of different parts of low erucic Brassica napus WY20’s seed were analyzed. There was a significant difference in oil content in different parts of the seed. The oil content in the outer cotyledon was the highest while embryonic axis had the lowest oil content. At the same time, the fatty acid composition also showed significant difference in different parts of the seed. The ratio of C16:0, C18:2, and C20:0 in embryonic axis was significantly higher than that in cotyledon. The ratio of C16:0 in the embryonic axis was about twice more than that of the cotyledons. The ratio of C18:1 and C20:1 in cotyledons was significantly higher than that in embryonic axis. C18:0 had the lowest content in the outer cotyledon and no difference in the inner cotyledons and embryonic axis. C18:3 had the highest content in the outer cotyledons and no difference between inner cotyledons and embryonic axis. Transcriptome analysis was performed for the inner cotyledon, outer cotyledon and embryonic axis of the 34-day-old seed. A total of 7192 differentially expressed genes (DEGs) were identified after pairwise comparison of gene expression of the three parts. There were much fewer DEGs between cotyledons and more DEGs between cotyledon and embryonic axis. These DEGs were enriched in biological processes such as photosynthesis, fatty acid metabolism and chlorophyll metabolism. Gene function annotations revealed that there were 355 genes involved in lipid metabolism, especially in the de novo fatty acid biosynthesis in plastid. This study suggests that transcriptional regulation of key genes involved in oil biosynthesis results in different oil contents and fatty acid compositions in different parts of seed in Brassica napus.

Key words: Brassica napus, different seed parts, oil content, fatty acid composition, transcriptional regulation

图1

甘蓝型油菜WY20种子不同部位含油量和脂肪酸组成 A: 显微镜明视野下WY20种子部位; B: 种子不同部位含油量; C: 种子不同部位重量百分比; D: 种子不同部位脂肪酸组成; E: 整颗种子脂肪酸组成。"

图2

3个种子部位差异表达基因分布情况 A: 样本间PCA分析; B: IC、OC和EA差异表达基因数目及百分比; C:差异表达基因维恩图。"

图3

油菜胚的3个部位差异表达基因GO功能分布 A: IC和OC差异表达基因功能分布; B: IC和EA差异表达基因功能分布; C: OC和EA差异表达基因功能分布。"

表1

甘蓝型油菜皖油20种子不同部位油脂合成差异表达基因"

基因名
Gene ID		 注释描述
Annotation description		 蛋白家族缩写
Protein family abbreviations		 分组
Group
BnaA03g13590D Fatty acid desaturase 3 FAD3 EA-IC-OC
BnaC03g16520D Fatty acid desaturase 3 FAD3 EA-IC/EA-OC
BnaC04g14820D Fatty acid desaturase 3 FAD3 EA-IC/EA-OC
BnaC04g40760D Fatty acid desaturase 3 FAD3 EA-IC/EA-OC
BnaA04g17150D Fatty acid desaturase 3 FAD3 EA-IC/EA-OC
BnaAnng09250D Fatty acid desaturase 2 FAD2 EA-IC/EA-OC
BnaA02g11570D Hydroxysteroid dehydrogenase 1 OBO EA-IC-OC
BnaA03g23490D Hydroxysteroid dehydrogenase 1 OBO EA-IC-OC
BnaC03g27860D Hydroxysteroid dehydrogenase 1 OBO EA-IC-OC
BnaCnng57830D Hydroxysteroid dehydrogenase 1 OBO EA-IC
BnaA09g02110D Oleosin OBO EA-OC
BnaA03g20420D Stearoyl-acyl-carrier-protein desaturase protein SAD EA-IC/EA-OC
BnaA01g32860D Stearoyl-acyl-carrier-protein desaturase protein SAD EA-IC/EA-OC
BnaC03g24420D Stearoyl-acyl-carrier-protein desaturase protein SAD EA-IC/EA-OC
BnaC09g41580D Stearoyl-acyl-carrier-protein desaturase protein SAD EA-IC/EA-OC
BnaA10g18080D Stearoyl-acyl-carrier-protein desaturase protein SAD EA-IC/EA-OC
BnaA05g03490D Stearoyl-acyl-carrier-protein desaturase SAD EA-OC
BnaC04g03030D Stearoyl-acyl-carrier-protein desaturase SAD EA-OC
BnaC09g19280D 3-ketoacyl-acyl carrier protein synthase I KASI EA-IC/EA-OC
BnaA02g24400D 3-ketoacyl-acyl carrier protein synthase I KASI EA-IC/EA-OC
BnaA06g36060D 3-ketoacyl-acyl carrier protein synthase I KASI EA-IC/EA-OC
BnaC06g35760D 3-ketoacyl-acyl carrier protein synthase II KAS II EA-IC/EA-OC
BnaA07g31890D 3-ketoacyl-acyl carrier protein synthase II KAS II EA-IC/EA-OC
BnaA07g21940D 3-ketoacyl-acyl carrier protein synthase II KAS II EA-IC/EA-OC
BnaC06g22680D 3-ketoacyl-acyl carrier protein synthase II KAS II EA-IC
BnaA04g07120D Acyl-ACP thioesterase FATA EA-IC/EA-OC
BnaCnng41490D Acyl-ACP thioesterase FATA EA-IC/EA-OC
BnaCnng00070D FATA acyl-ACP thioesterase FATA FATA EA-IC/EA-OC
BnaA07g05070D FATA acyl-ACP thioesterase FATA FATA EA-IC/EA-OC
BnaC03g75820D Ketoacyl-ACP Reductase KAR EA-IC/EA-OC
BnaA02g13310D Beta-ketoacyl reductase KAR EA-IC/EA-OC
BnaA07g26670D Beta-ketoacyl reductase KAR EA-IC/EA-OC
BnaC06g28830D Beta-ketoacyl reductase KAR EA-IC/EA-OC
BnaC09g16320D Acyl carrier protein ACP EA-IC/EA-OC
BnaC09g03000D Acyl carrier protein ACP EA-IC/EA-OC
BnaA09g03610D Acyl carrier protein ACP EA-IC/EA-OC
BnaAnng23710D Acyl carrier protein ACP EA-OC
BnaC03g45040D Enoyl-ACP Reductase ENR EA-IC/EA-OC
BnaC07g04330D Enoyl-ACP Reductase ENR EA-IC/EA-OC
BnaA03g38220D Enoyl-ACP Reductase ENR EA-IC/EA-OC
BnaAnng02240D E2 component of pyruvate dehydrogenase complex DHLAT EA-IC/EA-OC
BnaC06g08280D E2 component of pyruvate dehydrogenase complex DHLAT EA-IC/EA-OC
BnaC07g23030D E2 component of pyruvate dehydrogenase complex DHLAT EA-IC
BnaA06g33300D E2 component of pyruvate dehydrogenase complex DHLAT EA-IC
BnaAnng22560D Chloroplasticacetyl coenzyme A carboxylase BCCP EA-IC/EA-OC
BnaC09g42420D Biotin carboxyl carrier protein 2 BCCP EA-IC/EA-OC
BnaA03g02830D Thioesterase superfamily protein HAD EA-IC/EA-OC
BnaA02g00390D Thioesterase superfamily protein HAD EA-IC
BnaA07g20920D Long chain acyl-CoA synthetase 9 LACS9 EA-IC/EA-OC
BnaC06g20910D Long chain acyl-CoA synthetase 9 LACS9 EA-IC
BnaA01g17630D E3 component of pyruvate dehydrogenase complex LPD EA-IC/EA-OC
BnaCnng75250D Acetyl-CoA carboxylase ACCase EA-IC
BnaA05g12180D Acyl-carrier-protein MCMT EA-IC/EA-OC

图4

油脂合成相关差异表达基因在油脂代谢通路中的比较分析 维恩图中阴影代表差异表达基因的分布。红色数字表示蛋白家族对应基因TPM值的平均值在IC和EA间的比值, 绿色数字表示蛋白家族对应基因TPM值的平均值在OC和EA间的比值, 蓝色数字表示蛋白家族对应基因TPM值的平均值在IC和OC间的比值。编码蛋白质的基因: DHLAT: 二氢硫辛酰胺乙酰转移酶; LPD: 二氢硫辛酰胺脱氢酶; BCCP: 生物素羧基载体蛋白; ACCase: 乙酰辅酶a羧化酶; MCMT: 酰基载体蛋白; KASI: 3-酮酰基-酰基载体蛋白合酶I; KASII: 3-酮酰基-酰基载体蛋白合酶II; KASIII: 3-酮酰基-酰基载体蛋白合酶III; KAR: 酮脂酰还原酶; HAD: 硫酯酶蛋白; ENR: 烯酰ACP还原酶; ACP: 酰基载体蛋白; SAD: 硬脂酰-酰基载体蛋白脱饱和酶蛋白; FATA: FATA硫酯酶; FATB: FATB硫酯酶; LACS9: 长链酰基辅酶9; FAD2: 脂肪酸去饱和酶2; FAD3: 脂肪酸去饱和酶3; CPT: CDP胆碱-甘油二酯胆碱酯酶; PDCT: 磷脂酰胆碱-甘油二酯胆碱酯酶; GPAT: 磷酸甘油脂酰转移酶; PDAT: 磷脂-二酰甘油酰基转移酶; LPAAT: 溶血磷脂酸酰基转移酶; PAP: 磷脂酸磷酸酶; DGAT: 二酰甘油酰基转移酶; OBO: 油体蛋白; LTP: 脂质转运蛋白; ER: 内质网。"

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