园艺学报 ›› 2022, Vol. 49 ›› Issue (5): 958-972.doi: 10.16420/j.issn.0513-353x.2021-0103
梁沁1,2, 张延晖3, 康开权4, 刘瑾航1, 李亮1, 冯宇1, 王超1, 杨超1,*(), 李永裕1,*(
)
收稿日期:
2022-01-11
修回日期:
2022-04-01
出版日期:
2022-05-25
发布日期:
2022-05-25
通讯作者:
杨超,李永裕
E-mail:yycc32@126.com;lilin3182@163.com
基金资助:
LIANG Qin1,2, ZHANG Yanhui3, KANG Kaiquan4, LIU Jinhang1, LI Liang1, FENG Yu1, WANG Chao1, YANG Chao1,*(), LI Yongyu1,*(
)
Received:
2022-01-11
Revised:
2022-04-01
Online:
2022-05-25
Published:
2022-05-25
Contact:
YANG Chao,LI Yongyu
E-mail:yycc32@126.com;lilin3182@163.com
摘要:
为了探究miR168家族的进化特性及其在梨休眠进程中的表达模式,通过检索实验室前期构建的砂梨(‘黄花’梨)miRNA文库、转录组数据库和miRBase数据库,得到植物miR168家族成员的成熟体和前体序列。从‘黄花’梨花芽中克隆了两条miR168前体,发现梨miR168家族包含2条前体序列和4条成熟体序列。采用生物信息分析和实时荧光定量PCR(qRT-PCR),对梨miR168家族的成熟体序列及前体(pre-miR168)序列进行分析,与已知植物miR168构建系统发育进化树,预测其靶基因,分析其在梨休眠发生进程中的表达情况。结果表明,梨pre-miR168序列均能形成稳定的发夹结构;植物pre-miR168两端序列较保守,而中间序列不保守;mi168成熟体保守性较高,进化树分析表明梨pre-miR168与苹果的亲缘关系最近,miR168成熟体主要分为两大类。梨miR168调控的潜在靶基因主要包含Argonaute1蛋白基因(protein argonaute 1,AGO1)和胼胝质合成酶3基因(callose synthase 3,CalS3)共21条mRNA,都是以裂解靶标的方式进行调控。qRT-PCR分析结果表明,梨miR168a相对表达量在整个休眠期间呈先上升后下降的单峰动态变化;靶基因AGO1的表达呈上升趋势,靶基因CalS3的表达呈下降趋势;AGO1的表达趋势与梨花芽的萌芽率呈正相关关系,随着萌芽率的升高而升高,而CalS3表达趋势与梨花芽的萌芽率呈负相关关系,随萌芽率升高而降低。梨miR168与其他植物miR168的序列具有高度保守性,miR168与AGO1可能通过ABA来调控梨花芽的冬季休眠,通过上调表达梨miR168a而抑制CalS3表达的方式促进梨花芽冬季休眠的解除。
中图分类号:
梁沁, 张延晖, 康开权, 刘瑾航, 李亮, 冯宇, 王超, 杨超, 李永裕. miR168家族进化特性及其在砂梨休眠期的表达模式分析[J]. 园艺学报, 2022, 49(5): 958-972.
LIANG Qin, ZHANG Yanhui, KANG Kaiquan, LIU Jinhang, LI Liang, FENG Yu, WANG Chao, YANG Chao, LI Yongyu. Molecular Evolution of MiR168 Family and Their Expression Profiling During Dormancy of Pyrus pyrifolia[J]. Acta Horticulturae Sinica, 2022, 49(5): 958-972.
用途Purpose | 引物名称Primer name | 引物序列(5′-3′)Primer sequence |
---|---|---|
克隆Clone | ppy-miR168a-F | ATAGAGGCGTAGGCCTGTGT |
ppy-miR168a-R | TGATGGTGGGAATTGGGATAT | |
克隆Clone | ppy-miR168b-F | TGTCTGTTTGGGCGTTGATT |
ppy-miR168b-R | AAACTATTGTAGATCAGACAACAACGC | |
qRT-PCR | Ppy-miR168a | TCGCTTGGTGCAGGTCG |
AGO1-F | CACCAGCACCACAACAATA | |
AGO1-R | ACCACCACCATAGCCTCT | |
CalS3-F | AGAGGCGGATAACTCG | |
CalS3-R | GCAACACGAAGAATGG | |
Actin-F | CCATCCAGGCTGTTCTCTC | |
Actin-R | GCAAGGTCCAGACGAAGG | |
5SrRNA-F | TAGGCGTAGAGGAACCACACCAAT | |
5SrRNA-R | ACAGTATCGTCACCGCAGTAGAGT |
表1 本试验中所用引物
Table 1 The primers used in this experiment
用途Purpose | 引物名称Primer name | 引物序列(5′-3′)Primer sequence |
---|---|---|
克隆Clone | ppy-miR168a-F | ATAGAGGCGTAGGCCTGTGT |
ppy-miR168a-R | TGATGGTGGGAATTGGGATAT | |
克隆Clone | ppy-miR168b-F | TGTCTGTTTGGGCGTTGATT |
ppy-miR168b-R | AAACTATTGTAGATCAGACAACAACGC | |
qRT-PCR | Ppy-miR168a | TCGCTTGGTGCAGGTCG |
AGO1-F | CACCAGCACCACAACAATA | |
AGO1-R | ACCACCACCATAGCCTCT | |
CalS3-F | AGAGGCGGATAACTCG | |
CalS3-R | GCAACACGAAGAATGG | |
Actin-F | CCATCCAGGCTGTTCTCTC | |
Actin-R | GCAAGGTCCAGACGAAGG | |
5SrRNA-F | TAGGCGTAGAGGAACCACACCAAT | |
5SrRNA-R | ACAGTATCGTCACCGCAGTAGAGT |
科 Family | 物种 Species | 简写 Logogram | miR168数量 Quantity of miR168 | pre-miR168数量 Quantity of pre-miR168 |
---|---|---|---|---|
十字花科Brassicaceae | 拟南芥Arabidopsis thaliana | ath | 4 | 2 |
油菜Brassica napus | bna | 2 | 2 | |
琴叶拟南芥Arabidopsis lyrata | aly | 4 | 2 | |
芜菁Brassica rapa | bra | 6 | 3 | |
亚麻荠Camelina sativa | csa | 1 | 1 | |
禾本科Poaceae | 青稞Hordeum vulgare | hvu | 2 | 1 |
二穗短柄草Brachypodium distachyon | bdi | 2 | 1 | |
甘蔗属Saccharum sp | ssp | 1 | 1 | |
甘蔗Saccharum officinarum | sof | 2 | 2 | |
水稻Oryza sativa | osa | 3 | 2 | |
高粱Sorghum bicolor | sbi | 1 | 1 | |
玉米Zea mays | zma | 4 | 2 | |
山羊草Aegilops tauschii | ata | 1 | 1 | |
豆科Leguminosae | 大豆Glycine max | gma | 2 | 2 |
蒺藜苜蓿Medicago truncatula | mtr | 4 | 3 | |
大叶相思Acacia auriculiformis | aau | 1 | 1 | |
豇豆Vigna unguiculata | vun | 1 | 1 | |
百脉根Lotus japonicus | lja | 1 | 1 | |
科 Family | 物种 Species | 简写 Logogram | miR168数量 Quantity of miR168 | pre-miR168数量 Quantity of pre-miR168 |
杨柳科Salicaceae Mirb | 毛果杨Populus trichocarpa | ptc | 4 | 2 |
葡萄科Vitaceae Juss | 葡萄Vitis vinifera | vvi | 1 | 1 |
毛茛科Ranunculaceae Juss | 耧斗菜Aquilegia viridiflora | aqc | 1 | 1 |
芸香科Rutaceae Juss | 脐橙Citrus sinensis | csi | 2 | 1 |
柑橘Citrus reticulata | cre | 1 | 1 | |
克萊门柚Citrus clementina | ccl | 1 | 1 | |
大戟科Rosaceae Juss | 木薯Manihot esculenta | mes | 1 | 1 |
三叶橡胶树 Hevea brasiliensis | hbr | 1 | 1 | |
蓖麻 Ricinus communis | rco | 1 | 1 | |
葫芦科Cucurbitaceae | 甜瓜Cucumis melo | cme | 1 | 1 |
菊科Asteraceae Bercht | 刺苞菜蓟Cynara cardunculus | cca | 1 | 1 |
亚麻科Linaceae | 亚麻Linum usitatissimum | lus | 2 | 2 |
茄科Solanaceae Juss | 烟草Nicotiana tabacum | nta | 5 | 5 |
番茄Lycopersicon esculentum | sly | 4 | 2 | |
蔷薇科Rosaceae Juss | 苹果Malus domestica | mdm | 2 | 2 |
碧桃Prunus persica | ppe | 1 | 1 | |
森林草莓Fragaria vesca | fve | 1 | 1 | |
凤梨科Bromeliaceae Juss | 莺歌凤梨Vriesea carinata | vcr | 4 | 2 |
松科Pinaceae Lindl | 挪威云杉Picea abies | pab | 2 | 2 |
百合科Liliaceae Juss | 芦笋Asparagus officinalis | aof | 2 | 2 |
梧桐科Sterculiaceae | 可可Theobroma cacao | tcc | 1 | 1 |
表2 植物 miR168家族成员分布与数量
Table 2 Distribution and quantity of plant miR168 family members
科 Family | 物种 Species | 简写 Logogram | miR168数量 Quantity of miR168 | pre-miR168数量 Quantity of pre-miR168 |
---|---|---|---|---|
十字花科Brassicaceae | 拟南芥Arabidopsis thaliana | ath | 4 | 2 |
油菜Brassica napus | bna | 2 | 2 | |
琴叶拟南芥Arabidopsis lyrata | aly | 4 | 2 | |
芜菁Brassica rapa | bra | 6 | 3 | |
亚麻荠Camelina sativa | csa | 1 | 1 | |
禾本科Poaceae | 青稞Hordeum vulgare | hvu | 2 | 1 |
二穗短柄草Brachypodium distachyon | bdi | 2 | 1 | |
甘蔗属Saccharum sp | ssp | 1 | 1 | |
甘蔗Saccharum officinarum | sof | 2 | 2 | |
水稻Oryza sativa | osa | 3 | 2 | |
高粱Sorghum bicolor | sbi | 1 | 1 | |
玉米Zea mays | zma | 4 | 2 | |
山羊草Aegilops tauschii | ata | 1 | 1 | |
豆科Leguminosae | 大豆Glycine max | gma | 2 | 2 |
蒺藜苜蓿Medicago truncatula | mtr | 4 | 3 | |
大叶相思Acacia auriculiformis | aau | 1 | 1 | |
豇豆Vigna unguiculata | vun | 1 | 1 | |
百脉根Lotus japonicus | lja | 1 | 1 | |
科 Family | 物种 Species | 简写 Logogram | miR168数量 Quantity of miR168 | pre-miR168数量 Quantity of pre-miR168 |
杨柳科Salicaceae Mirb | 毛果杨Populus trichocarpa | ptc | 4 | 2 |
葡萄科Vitaceae Juss | 葡萄Vitis vinifera | vvi | 1 | 1 |
毛茛科Ranunculaceae Juss | 耧斗菜Aquilegia viridiflora | aqc | 1 | 1 |
芸香科Rutaceae Juss | 脐橙Citrus sinensis | csi | 2 | 1 |
柑橘Citrus reticulata | cre | 1 | 1 | |
克萊门柚Citrus clementina | ccl | 1 | 1 | |
大戟科Rosaceae Juss | 木薯Manihot esculenta | mes | 1 | 1 |
三叶橡胶树 Hevea brasiliensis | hbr | 1 | 1 | |
蓖麻 Ricinus communis | rco | 1 | 1 | |
葫芦科Cucurbitaceae | 甜瓜Cucumis melo | cme | 1 | 1 |
菊科Asteraceae Bercht | 刺苞菜蓟Cynara cardunculus | cca | 1 | 1 |
亚麻科Linaceae | 亚麻Linum usitatissimum | lus | 2 | 2 |
茄科Solanaceae Juss | 烟草Nicotiana tabacum | nta | 5 | 5 |
番茄Lycopersicon esculentum | sly | 4 | 2 | |
蔷薇科Rosaceae Juss | 苹果Malus domestica | mdm | 2 | 2 |
碧桃Prunus persica | ppe | 1 | 1 | |
森林草莓Fragaria vesca | fve | 1 | 1 | |
凤梨科Bromeliaceae Juss | 莺歌凤梨Vriesea carinata | vcr | 4 | 2 |
松科Pinaceae Lindl | 挪威云杉Picea abies | pab | 2 | 2 |
百合科Liliaceae Juss | 芦笋Asparagus officinalis | aof | 2 | 2 |
梧桐科Sterculiaceae | 可可Theobroma cacao | tcc | 1 | 1 |
图3 植物miR168成熟体进化树(a)及序列多重比较(b) ath. 拟南芥;ptc. 毛果杨;vvi. 葡萄;crt. 柑橘;csi. 甜橙;mdm. 苹果;ppe. 碧桃;fve. 野草莓。
Fig. 3 Evolutionary tree of miR168(a)and multiple sequences comparison(b) ath. Arabidopsis thaliana;ptc. Populus trichocarpa;vvi. Vitis vinifera;crt. Citrus reticulata;csi. Citrus sinensis;mdm. Malus × domestica;ppe. Prunus persica;fve. Fragaria vesca.
图5 梨miR168前体二级茎环结构 颜色越偏红色,碱基处于配对的概率越大,反之越小。
Fig. 5 Analysis of secondary stem-loop structure of Pyrus pyrifolia miR168 precursors. The redder the color,the more likely the base is to be in a pair,and vice versa.
靶基因注释 Target gene annotation | 靶基因编号 Target accession | 期望值 Expection |
---|---|---|
AGO1蛋白 Protein argonaute 1-like transcript variant | XM_018645622.1 | 0 |
XM_018647395.1 | 0 | |
XM_009349800.2 | 0 | |
XM_018645623.1 | 0 | |
XM_018647396.1 | 0 | |
XM_009357682.2 | 0 | |
胼胝质合成酶3(CalS3) Callose synthase 3(CalS3)transcript variant | XM_018649103.1 | 2.5 |
XM_009376827.2 | 2.5 | |
XM_018651590.1 | 2.5 | |
XM_018651591.1 | 2.5 | |
XM_009365800.2 | 2.5 | |
XM_018649104.1 | 2.5 | |
XM_009365801.2 | 2.5 | |
XM_009365802.2 | 2.5 | |
XM_009365803.2 | 2.5 | |
AGO1蛋白Protein argonaute 1-like transcript variant | XM_009359066.2 | 2.5 |
XM_009349798.2 | 2.5 | |
XM_009359065.2 | 2.5 | |
XM_009349797.2 | 2.5 | |
XM_009357684.2 | 2.5 | |
XM_009357683.2 | 2.5 |
表3 梨miR168家族成员(ppy-miR168a-5p,ppy-miR168,ppy-miR168b)靶基因预测
Table 3 Target genes prediction of pear miR168 family(ppy-miR168a-5p,ppy-miR168,ppy-miR168b)
靶基因注释 Target gene annotation | 靶基因编号 Target accession | 期望值 Expection |
---|---|---|
AGO1蛋白 Protein argonaute 1-like transcript variant | XM_018645622.1 | 0 |
XM_018647395.1 | 0 | |
XM_009349800.2 | 0 | |
XM_018645623.1 | 0 | |
XM_018647396.1 | 0 | |
XM_009357682.2 | 0 | |
胼胝质合成酶3(CalS3) Callose synthase 3(CalS3)transcript variant | XM_018649103.1 | 2.5 |
XM_009376827.2 | 2.5 | |
XM_018651590.1 | 2.5 | |
XM_018651591.1 | 2.5 | |
XM_009365800.2 | 2.5 | |
XM_018649104.1 | 2.5 | |
XM_009365801.2 | 2.5 | |
XM_009365802.2 | 2.5 | |
XM_009365803.2 | 2.5 | |
AGO1蛋白Protein argonaute 1-like transcript variant | XM_009359066.2 | 2.5 |
XM_009349798.2 | 2.5 | |
XM_009359065.2 | 2.5 | |
XM_009349797.2 | 2.5 | |
XM_009357684.2 | 2.5 | |
XM_009357683.2 | 2.5 |
处理日期(M-D)Treatment date | ppy-miR168a | AGO1 | CalS3 |
---|---|---|---|
11-01 | 0.22 | 0 | 4.00 |
11-25 | 3.37 | 0.05 | 1.03 |
12-27 | 1.12 | 0.58 | 0.75 |
01-23 | 1.00 | 5.91 | 0.59 |
相关系数Correlation coefficient | -0.223 | -0.503 |
表4 ppy-miR168a及其靶基因表达量相关性分析
Table 4 Correlation analysis of ppy-miR168a and its target genes expression
处理日期(M-D)Treatment date | ppy-miR168a | AGO1 | CalS3 |
---|---|---|---|
11-01 | 0.22 | 0 | 4.00 |
11-25 | 3.37 | 0.05 | 1.03 |
12-27 | 1.12 | 0.58 | 0.75 |
01-23 | 1.00 | 5.91 | 0.59 |
相关系数Correlation coefficient | -0.223 | -0.503 |
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