园艺学报 ›› 2022, Vol. 49 ›› Issue (2): 352-364.doi: 10.16420/j.issn.0513-353x.2020-1013
收稿日期:
2021-07-01
修回日期:
2021-12-23
出版日期:
2022-02-25
发布日期:
2022-02-28
通讯作者:
孙黎
E-mail:sunlishz@126.com
基金资助:
ZHOU Zhiming, YANG Jiabao, ZHANG Cheng, ZENG Linglu, MENG Wanqiu, SUN Li()
Received:
2021-07-01
Revised:
2021-12-23
Online:
2022-02-25
Published:
2022-02-28
Contact:
SUN Li
E-mail:sunlishz@126.com
摘要:
利用生物信息学技术从向日葵(Helianthus annuus)基因组中鉴定了13个长链酰基辅酶A合成酶(long-chain acyl-CoA synthases,LACSs)家族基因,其分布在向日葵8条染色体上。系统发育树显示,HaLACS家族成员分为6个亚族,且同一亚族成员具有相似的基因结构和蛋白保守基序。基因复制分析发现HaLACS家族扩增的主要因素是片段复制。HaLACS的启动子区域富含逆境响应和植物激素调控相关元件。表达谱分析显示HaLACS在向日葵不同组织中的表达存在差异。比较2个向日葵品种(高/低亚油酸含量)中HaLACS在种子发育不同阶段的表达,HaLACS呈现差异表达。qRT-PCR结果表明,200 mmol · L-1 NaCl处理24 h后,7个HaLACS在茎中的表达量与同期对照相比显著上调;100 μmol · L-1 ABA 处理3 h后,8个HaLACS在根中的表达量显著上调;15% PEG 6000模拟干旱胁迫后,多数HaLACS在不同组织中呈现诱导表达。综上,向日葵LACS基因家族不仅参与了向日葵脂质的合成,而且在应对非生物胁迫时发挥重要作用。
中图分类号:
周至铭, 杨佳宝, 张程, 曾令露, 孟晚秋, 孙黎. 向日葵LACS家族鉴定及响应非生物胁迫表达分析[J]. 园艺学报, 2022, 49(2): 352-364.
ZHOU Zhiming, YANG Jiabao, ZHANG Cheng, ZENG Linglu, MENG Wanqiu, SUN Li. Genome-wide Identification and Expression Analyses of Long-chain Acyl-CoA Synthetases Under Abiotic Stresses in Helianthus annuus[J]. Acta Horticulturae Sinica, 2022, 49(2): 352-364.
基因Gene | 正向引物(5′-3′)Forward primer | 反向引物(5′-3′)Reverse primer |
---|---|---|
18S rRNA | CTACCACATCCAAGGAAGGCAG | CGACAGAAGGGACGAGTAAACC |
HaLACS1-1 | GAGCTCGAAAACGGAACGTG | ATTGCAAGCCTCCATCGCTA |
HaLACS1-2 | TTGGACCATGCCGAAGTTGA | AAGGAACTCGTTCCACGCAT |
HaLACS2 | AGAAACAGACAAACCGGGCA | ACGCCTTCTTTACAGTGCCT |
HaLACS4-1 | GGTTATGAGCATGGAGGCGT | CCACGCCAAAATTTCCGACT |
HaLACS4-2 | GGTTATGAGCATGGAGGCGT | GGTTATGAGCATGGAGGCGT |
HaLACS4-3 | CCTCCGATTGACGGTTTGGA | CTCAACTCCACAAGCTCGGA |
HaLACS4-4 | CGGTGAAAGACGGAACTCCA | CGACGCCACATGAACGGATA |
HaLACS4-5 | GAAAACAATCGTGAGCTTTGGGA | AGTTCAACGATACTTTCGCGCT |
HaLACS6 | CACTGCCGGAGAGTTCTTCA | TGCTTCCCCGTAAGTTGTCC |
HaLACS7 | TCATCTTCTTCCCTCCACCGA | ATCTGGTGACAAGCTTCAGCG |
HaLACS8 | GCAGATGGATGTTGGGGGAT | CGCCAACTTCAACTGGAACG |
HaLACS9-1 | ACTCTCGCCCTTCGGAAATC | TTTCAAATGCCCTCCCGTCT |
HaLACS9-2 | ATTGCATTGCAGGCGTGTTT | TTTCCAACTGCTGCTTCCGT |
表1 qRT-PCR引物序列
Table 1 Primers used for qRT-PCR
基因Gene | 正向引物(5′-3′)Forward primer | 反向引物(5′-3′)Reverse primer |
---|---|---|
18S rRNA | CTACCACATCCAAGGAAGGCAG | CGACAGAAGGGACGAGTAAACC |
HaLACS1-1 | GAGCTCGAAAACGGAACGTG | ATTGCAAGCCTCCATCGCTA |
HaLACS1-2 | TTGGACCATGCCGAAGTTGA | AAGGAACTCGTTCCACGCAT |
HaLACS2 | AGAAACAGACAAACCGGGCA | ACGCCTTCTTTACAGTGCCT |
HaLACS4-1 | GGTTATGAGCATGGAGGCGT | CCACGCCAAAATTTCCGACT |
HaLACS4-2 | GGTTATGAGCATGGAGGCGT | GGTTATGAGCATGGAGGCGT |
HaLACS4-3 | CCTCCGATTGACGGTTTGGA | CTCAACTCCACAAGCTCGGA |
HaLACS4-4 | CGGTGAAAGACGGAACTCCA | CGACGCCACATGAACGGATA |
HaLACS4-5 | GAAAACAATCGTGAGCTTTGGGA | AGTTCAACGATACTTTCGCGCT |
HaLACS6 | CACTGCCGGAGAGTTCTTCA | TGCTTCCCCGTAAGTTGTCC |
HaLACS7 | TCATCTTCTTCCCTCCACCGA | ATCTGGTGACAAGCTTCAGCG |
HaLACS8 | GCAGATGGATGTTGGGGGAT | CGCCAACTTCAACTGGAACG |
HaLACS9-1 | ACTCTCGCCCTTCGGAAATC | TTTCAAATGCCCTCCCGTCT |
HaLACS9-2 | ATTGCATTGCAGGCGTGTTT | TTTCCAACTGCTGCTTCCGT |
转录本ID Transcript ID | 基因名称 Gene name | CDS/bp | 大小/aa Size | 蛋白分子量/kD Molecular weight | 等电点 pI | 拟南芥同源基因 Arabidopsis homologous gene | |
---|---|---|---|---|---|---|---|
登录号Accession No. | 名称Name | ||||||
OTG27997 | HaLACS1-1 | 1 980 | 659 | 74.4 | 5.82 | AT2G47240 | AtLACS1 |
OTF93791 | HaLACS1-2 | 1 986 | 661 | 74.8 | 7.23 | AT2G47240 | AtLACS1 |
OTG12244 | HaLACS2 | 1 980 | 659 | 73.5 | 5.70 | AT1G49430 | AtLACS2 |
OTG32156 | HaLACS4-1 | 1 986 | 661 | 73.5 | 6.07 | AT4G23850 | AtLACS4 |
OTG33063 | HaLACS4-2 | 1 995 | 664 | 74.0 | 6.69 | AT4G23850 | AtLACS4 |
OTG20245 | HaLACS4-3 | 1 980 | 659 | 73.5 | 6.50 | AT4G23850 | AtLACS4 |
OTG02621 | HaLACS4-4 | 1 974 | 657 | 73.4 | 6.69 | AT4G23850 | AtLACS4 |
OTG02629 | HaLACS4-5 | 1 983 | 660 | 73.8 | 8.49 | AT4G23850 | AtLACS4 |
OTG11238 | HaLACS6 | 2 094 | 697 | 76.3 | 7.51 | AT3G05970 | AtLACS6 |
OTG31091 | HaLACS7 | 2 097 | 698 | 77.0 | 6.58 | AT5G27600 | AtLACS7 |
OTG29824 | HaLACS8 | 2 163 | 720 | 78.6 | 7.86 | AT2G04350 | AtLACS8 |
OTG15459 | HaLACS9-1 | 2 091 | 696 | 76.2 | 6.10 | AT1G77590 | AtLACS9 |
OTG07054 | HaLACS9-2 | 2 121 | 706 | 77.4 | 6.78 | AT1G77590 | AtLACS9 |
表2 向日葵LACS基因基本信息
Table 2 Detailed information of LACS genes in sunflower genome
转录本ID Transcript ID | 基因名称 Gene name | CDS/bp | 大小/aa Size | 蛋白分子量/kD Molecular weight | 等电点 pI | 拟南芥同源基因 Arabidopsis homologous gene | |
---|---|---|---|---|---|---|---|
登录号Accession No. | 名称Name | ||||||
OTG27997 | HaLACS1-1 | 1 980 | 659 | 74.4 | 5.82 | AT2G47240 | AtLACS1 |
OTF93791 | HaLACS1-2 | 1 986 | 661 | 74.8 | 7.23 | AT2G47240 | AtLACS1 |
OTG12244 | HaLACS2 | 1 980 | 659 | 73.5 | 5.70 | AT1G49430 | AtLACS2 |
OTG32156 | HaLACS4-1 | 1 986 | 661 | 73.5 | 6.07 | AT4G23850 | AtLACS4 |
OTG33063 | HaLACS4-2 | 1 995 | 664 | 74.0 | 6.69 | AT4G23850 | AtLACS4 |
OTG20245 | HaLACS4-3 | 1 980 | 659 | 73.5 | 6.50 | AT4G23850 | AtLACS4 |
OTG02621 | HaLACS4-4 | 1 974 | 657 | 73.4 | 6.69 | AT4G23850 | AtLACS4 |
OTG02629 | HaLACS4-5 | 1 983 | 660 | 73.8 | 8.49 | AT4G23850 | AtLACS4 |
OTG11238 | HaLACS6 | 2 094 | 697 | 76.3 | 7.51 | AT3G05970 | AtLACS6 |
OTG31091 | HaLACS7 | 2 097 | 698 | 77.0 | 6.58 | AT5G27600 | AtLACS7 |
OTG29824 | HaLACS8 | 2 163 | 720 | 78.6 | 7.86 | AT2G04350 | AtLACS8 |
OTG15459 | HaLACS9-1 | 2 091 | 696 | 76.2 | 6.10 | AT1G77590 | AtLACS9 |
OTG07054 | HaLACS9-2 | 2 121 | 706 | 77.4 | 6.78 | AT1G77590 | AtLACS9 |
重复基因对 Duplicated gene pairs | 非同义替 换率Ka | 同义替换率 Ks | Ka/Ks | 重复类型 Duplicated type | 选择压力 Selective pressure | 分化时间/ Mya Divergence time |
---|---|---|---|---|---|---|
HaLACS9-1/HaLACS9-2 | 0.0540 | 0.5316 | 0.1016 | 片段重复Segmental | 纯化Purifying | 17.72 |
HaLACS4-1/HaLACS4-5 | 0.0739 | 0.6713 | 0.1101 | 片段重复Segmental | 纯化Purifying | 22.38 |
HaLACS4-1/HaLACS4-4 | 0.0905 | 0.6463 | 0.1400 | 片段重复Segmental | 纯化Purifying | 21.54 |
HaLACS4-1/HaLACS4-3 | 0.1003 | 1.0273 | 0.0976 | 片段重复Segmental | 纯化Purifying | 34.24 |
HaLACS4-4/HaLACS4-5 | 0.0600 | 0.3516 | 0.1706 | 串联重复Tandem | 纯化Purifying | 11.72 |
表3 向日葵LACS家族成员基因复制事件和发散时间分析
Table 3 Gene duplication events and divergence time of the LACS genes in Helianthus annuus
重复基因对 Duplicated gene pairs | 非同义替 换率Ka | 同义替换率 Ks | Ka/Ks | 重复类型 Duplicated type | 选择压力 Selective pressure | 分化时间/ Mya Divergence time |
---|---|---|---|---|---|---|
HaLACS9-1/HaLACS9-2 | 0.0540 | 0.5316 | 0.1016 | 片段重复Segmental | 纯化Purifying | 17.72 |
HaLACS4-1/HaLACS4-5 | 0.0739 | 0.6713 | 0.1101 | 片段重复Segmental | 纯化Purifying | 22.38 |
HaLACS4-1/HaLACS4-4 | 0.0905 | 0.6463 | 0.1400 | 片段重复Segmental | 纯化Purifying | 21.54 |
HaLACS4-1/HaLACS4-3 | 0.1003 | 1.0273 | 0.0976 | 片段重复Segmental | 纯化Purifying | 34.24 |
HaLACS4-4/HaLACS4-5 | 0.0600 | 0.3516 | 0.1706 | 串联重复Tandem | 纯化Purifying | 11.72 |
图2 向日葵和其他物种LACS基因家族系统进化分析 绿色圆圈代表单子叶植物,红色圆圈代表双子叶植物。 Ec:大肠杆菌(原核生物);Sc:酿酒酵母(真菌);Pt:三角褐指藻(藻类);Pp:小立碗藓(藓);Sm:江南卷柏(蕨);Mp:地钱(苔);At:拟南芥;Ha:向日葵;Ls:莴苣;Eg:油棕;Zm:玉米;Os:水稻;Hs:人;Mm:家鼠。
Fig. 2 Phylogenetic analysis of LACS gene family in sunflower and other species The green circles indicate monocotyledonous plants,the red circle indicates a dicot plant. Ec:Escherichia coli(prokaryote);Sc:Saccharomyces cerevisiae(fungi);Pt:Phaeodac tylumtricornutum(algae);Pp:Physcomitrella patens(moss);Sm:Selaginella moellendorffii(fern);Mp:Marchantia polymorpha(moss);At:Arabidopsis thaliana;Ha:Helianthus annuus;Ls:Lactuca sativa;Eg:Elaeis guineensis;Zm:Zea mays;Os:Oryza sativa;Hs:Homo sapiens;Mm:Mus musculus.
图4 向日葵HaLACS与其他同源蛋白的氨基酸多序列比对 Ec:大肠杆菌;At:拟南芥;Sc:酿酒酵母;Ha:向日葵。
Fig. 4 Multiple sequence alignment of the HaLACS amino acid sequences with other proteins Ec:Escherichia coli;At:Arabidopsis thaliana;Sc:Selaginella moellendorffii;Ha:Helianthus annuus.
图7 HaLACS在2个向日葵品系不同发育阶段种子(A)和不同组织(B)的表达模式
Fig. 7 Expression levels in different seed development stages(A)of two sunflower cultivars and different tissues(B)
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