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作物学报 ›› 2022, Vol. 48 ›› Issue (11): 2934-2944.doi: 10.3724/SP.J.1006.2022.14243

• 研究简报 • 上一篇    下一篇

结球甘蓝类钙调蛋白CMLs与花粉萌发NPG1及NPGRs相互作用研究

许彬1,2(), 曹绍玉1,2(), 苏甜1, 彭梦玲1,2, 吕霞1, 李振林1, 张国平1, 许俊强1,*()   

  1. 1云南农业大学云南省滇台特色农业产业化工程研究中心, 云南昆明 650201
    2云南农业大学园林园艺学院, 云南昆明 650201
  • 收稿日期:2021-12-22 接受日期:2022-03-25 出版日期:2022-11-12 网络出版日期:2022-04-20
  • 通讯作者: 许俊强
  • 作者简介:第一作者联系方式: 许彬, E-mail: 1650379536@qq.com;
    曹绍玉, E-mail: 1194642629@qq.com第一联系人：** 同等贡献。
  • 基金资助:
    本研究由云南省重大科技计划项目(202102AE09005);云南省科技人才和平台计划项目(202205AF150017);云南省科技人才和平台计划项目(202005AF150035);云南省科技人才和平台计划项目(2019IC009);国家自然科学基金项目(31560560);昆明市科技计划项目(2019-1-H-24615)

Interactions between CMLs and NPG1 and related proteins in pollen germination of Brassica oleracea L. var. capitata

XU Bin1,2(), CAO Shao-Yu1,2(), SU Tian1, PENG Meng-Ling1,2, LYU Xia1, LI Zhen-Lin1, ZHANG Guo-Ping1, XU Jun-Qiang1,*()   

  1. 1Dian-Tai Engineering Research Center for Characteristic Agriculture Industrialization of Yunnan Province, Yunnan Agricultural University, Kunming 650201, Yunnan, China
    2College of Landscape and Horticulture, Yunnan Agricultural University, Kunming 650201, Yunnan, China
  • Received:2021-12-22 Accepted:2022-03-25 Published:2022-11-12 Published online:2022-04-20
  • Contact: XU Jun-Qiang
  • Supported by:
    The Major Projects of Yunnan Science and Technology Plan(202102AE09005);The Science and Technology Talents and Platform Plan of Yunnan Province(202205AF150017);The Science and Technology Talents and Platform Plan of Yunnan Province(202005AF150035);The Science and Technology Talents and Platform Plan of Yunnan Province(2019IC009);The National Natural Science Foundation of China(31560560);The Science and Technology Plan Project of Kunming(2019-1-H-24615)

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摘要：

类钙调蛋白作为钙调蛋白的家族之一, 调控植物生长发育、激素调控及各种胁迫, 对类钙调蛋白对参与花粉萌发及伸长的深入研究具有重要意义。为研究结球甘蓝花粉CMLs与NPG1及相关蛋白互作关系, 探索CMLs在花粉发育及萌发过程的功能, 本研究以结球甘蓝ZG为材料, 克隆到12个花粉发育及萌发相关CMLs基因, 及花粉萌发基因NPG1、NPGR1和NPGR2, 通过酵母双杂交试验检测编码蛋白相互之间的互作关系, 并检测NPG1、NPGR1和NPGR2在不同授粉时间后的表达水平。结果表明, 12个CMLs基因开放阅读框(open reading frame, ORF)在447~1095 bp之间, 其中CML20、CML48、CML49、CML50均含有4个内含子, CML21含有2个内含子, 其余均不含内含子; 编码蛋白均不含信号肽和跨膜域, 均为胞外蛋白, 且均为亲水性蛋白; 分别含有2~4个EF-Hands结构域, 氨基酸数目在148~364个氨基酸之间, 亚细胞定位预测主要定位于细胞膜、液泡及细胞核中; 结球甘蓝NPG1、NPGR1、NPGR2分别聚类到不同的进化枝上, 与油菜、白菜的亲缘关系最近, 与拟南芥亲缘关系较近; 酵母双杂交试验表明, CML13、CML21、CML24、CML42不能与NPG1、NPGR1及NPGR2在体外发生相互作用, CML15、CML20、CML25、CML39、CML41、CML48、CML49均能与NPG1、NPGR1及NPGR2在体外发生相互作用, CML50可与NPGR1、NPGR2互作, 而不能与NPG1发生互作。表达分析表明, NPG1表达量在授粉后的柱头中呈显著上升趋势, NPGR1、NPGR2表达量变化均不显著; CMLs在不同授粉时间均有不同的表达量变化, 其中CML15、CML20、CML39、CML41、CML49均呈现出先升高后下降的趋势。本研究在结球甘蓝中筛选得到能与NPG1、NPGRs互作的CMLs蛋白, 说明多个CMLs能参与花粉萌发及花粉管伸长过程, 期望为Ca2+信号系统参与花粉萌发及花粉管伸长的机制研究提供参考。

关键词: 结球甘蓝, 花粉, CMLs, NPG1, NPGRs, 蛋白互作

Abstract:

As one of calmodulin families, calmodulin-like (CMLs) regulate plant growth and development, hormone regulation, stresses, which have great significance to the further study of calmodulin-like protein involved in pollen germination and growth. In this study, to explore the interaction between CMLs and NPG1 related genes in pollen development of Brassica oleracea L. var. capitata, and to explore the function of CMLs in pollen development and germination, 12 CMLs genes related to pollen development and germination, and pollen germination genes NPG1, NPGR1, and NPGR2 were cloned from “ZG”. Interactions between them were detected by yeast two-hybrid test. and the relative expression levels of NPG1, NPGR1, and NPGR2 after different pollination time was detected. The results showed that the open reading frames (ORFs) of 12 CMLs genes ranged from 447 bp to 1095 bp, among which CML20, CML48, CML49, and CML50 contained 4 introns, CML21 contained 2 introns, and the others contained no introns. All of them contained no signal peptide and transmembrane domain, all of them were extracellular proteins and hydrophilic proteins which contain 2-4 EF-hands, with amino acids range of 148-364 aa. Subcellular localization was mainly located in cell membrane, vacuole, and nucleus. NPG1, NPGR1, and NPGR2 were clustered on different evolutionary branches, which were the closest to Brassica rape and Brassica napus, and closer to Arabidopsis thaliana. Yeast two-hybrid showed that CML13, CML21, CML24, CML42 could not interact with NPG1, NPGR1, and NPGR2 in vitro, CML15, CML20, CML25, CML39, CML41, CML48, and CML49 could interact with NPG1, NPGR1, and NPGR2, CML50 could interact with NPGR1 and NPGR2, but not with NPG1. The relative expression level of NPG1 increased significantly after pollination, while the relative expression levels of NPGR1 and NPGR2 didn’t change significantly. The relative expression levels of CMLs genes were varied at different pollination times, and CML15, CML20, CML39, CML41, and CML49 were a trend of first increasing and then decreasing. In conclusion, several CMLs proteins were screened which can interact with NPG1 and NPGRs, indicating that CMLs can participate in the process of pollen germination and pollen tube elongation, and it was expected to provide references for the research on the mechanism of Ca2+ signaling system participating in pollen germination and pollen tube elongation.

Key words: Brassica oleracea L. var. capitata, pollen, calmodulin-like protein (CMLs), NPG1, NPGRs, protein interaction

表1

本研究中用到的引物"

基因 Gene name 正向引物 Forward primer (5°-3°) 反向引物 Reverse primer (5°-3°)
CML13 GAGCGAAATCCGACCCGGCTT CTACTTAGCAACCATCCTGGCA
CML15 CTCTCATCTGTGTTTTGAGG TCATGAATTAATTTTTAAACCA
CML20 CGAGAAATCGACGCGGAAAGAGCT ATAACAGAGACCTCATCAGTACC
CML21 CGATCTGATCTCAAAGCAGGA CACAAACATCAAGCCTTGTCA
CML24 CATCGTCTTCTTCTCCACCACA CCTAAAATACGATCATCAAGC
CML25 GATCCAGAGATTCATTCGCTTCCTT GTGAAACTCTACCACTGACCAC
CML39 CATCATGAAGAACACTCAACGT CAACATTGGATGGGCTGATGCCTTA
CML41 CCAATATGGCAACTCCCCAAG CTTCATAAGCATTTCGAGCAA
CML42 GTCAAGGAAGAGGATAAATGG TGAAGTCAAGAAGGGATAACG
CML48 CTTGATGATGGGACTGGCAACACTG CATATGAGGCAATGAACGGAATGAC
CML49 ATGTCTGGTTACCCTCCTTCGAGC TTAAGCGACGAGGAACGGTAGAAC
CML50 TTCCCAGCGATGTCAGGTTACC GTTGTTGTTGTATGTTCGTCG
NPG1 GGCTCTGCATGAAAACAACTGA TTAAAGAATGGTTGAGAAGCT
NPGR1 ATGTTGTGTGCTTGTTCAGGCG TCAAATGAAACTCTGCACCGG
NPGR2 ATGAGGAACAGCGAGACTAGGC TCATCTGAATGGCTCCACTGGC
qCML15 CTCGGTCTCAAACCATCAGGAGA GTTACCGATCTCTTCGTTCAGG
qCML20 ATGTCGAGTCTATACAGAGGTG TGCTCTTCCGTCATCTCGAAACC
qCML25 GTTCGGCGTCTAGAACCGGAGCA TGGAACCTCGTGGCCTAGACTTG
qCML39 TCAAGACACTGGGAGAGCAA CAAAATCCAGCATCCCATCT
qCML41 GTCACCACTCCCATGACGAGCT TGCCTCGTGAGTTATATACTCAC
qCML48 TCAGCTTGATGATGGGACTG TAACCGGGGTCCTTCTCTTT
qCML49 GGAGACCACAAGCCGCACAAA CGCCTGGAAGCAAGTCACGAT
qCML50 ACCAACGCCATGAAGATAGG CATCTATTCTCCCGCTCCTG
qNPG1 TAGTCAGACCTCGGTGCTC GCTCCTCGCTGCAAACCTTA
qNPGR1 CGAATCAAGACAGATACCG CATAGCATACGCCAAGGAA
qNPGR2 AACTACGAGGAGGCGAGGGC GAGGCTGAGGAGGAGCAACA
β-Actin TATCAACTACCAGCCACC GAACACCTCAGCTACACTC

图1

结球甘蓝CMLs基因克隆 M: Trans2K plus II DNA marker; 1~12分别为CML13、CML15、CML20、CML21、CML24、CML25、CML39、CML41、CML42、CML48、CML49、CML50基因的克隆。"

表2

结球甘蓝CMLs基因基本信息"

基因
Gene name		 编码区
Coding
region (bp)		 内含子
Introns		 跨膜域
Transmembrane		 信号肽
Signal peptide		 预测的蛋白 Predicted protein 疏水性
Hydrophobicity		 亚细胞定位
Subcellular
localization
EF-
hand		 氨基酸
Amino acid (aa)		 分子量
Molecular weight (kD)		 pI
CML13 447 0 无No 无No 3 148 16.40 4.62 亲水
Hydrophilic		 细胞膜
Cell membrane
CML15 471 0 无No 无No 4 156 16.99 3.94 亲水
Hydrophilic		 液泡
Vacuole
CML20 510 4 无No 无No 4 169 19.60 4.55 亲水
Hydrophilic		 细胞膜
Cell membrane
CML21 678 2 无No 无No 4 225 25.96 4.17 亲水
Hydrophilic		 细胞膜
Cell membrane
CML24 447 0 无No 无No 4 158 17.40 4.64 亲水
Hydrophilic		 细胞膜, 液泡
Cell membrane, vacuole
CML25 552 0 无No 无No 4 183 20.14 4.42 亲水
Hydrophilic		 液泡
Vacuole
CML39 510 0 无No 无No 4 169 19.25 4.35 亲水
Hydrophilic		 液泡
Vacuole
CML41 612 0 无No 无No 4 203 22.64 4.97 亲水
Hydrophilic		 细胞膜
Cell membrane
CML42 567 0 无No 无No 3 188 20.70 4.25 亲水
Hydrophilic		 细胞膜, 液泡
Cell membrane, vacuole
CML48 672 4 无No 无No 2 223 25.28 4.53 亲水
Hydrophilic		 细胞核
Nucleus
CML49 954 4 无No 无No 2 317 33.51 6.93 亲水
Hydrophilic		 细胞核
Nucleus
CML50 1095 4 无No 无No 2 364 38.06 6.27 亲水
Hydrophilic		 细胞膜, 细胞核
Cell membrane, nucleus

图2

结球甘蓝CMLs的进化树和基因结构图 左侧为进化树, 右侧为甘蓝CMLs的基因结构图。"

表3

结球甘蓝NPG1、NPGR1及NPGR2基本信息"

基因
Gene		 编码区
Coding region
(bp)		 内含子
Introns		 跨膜域
Transmembrane		 信号肽
Signal peptide		 预测的蛋白 Predicted proteins 疏水性
Hydrophobicity		 亚细胞定位
Subcellular
localization
TPR 氨基酸
Amino
acid (aa)		 分子量Molecular
weight (kD)		 pI
NPG1 2043 4 无No 无No 3 680 75.31 5.98 亲水Hydrophilic 细胞核Nucleus
NPGR1 2109 2 无No 无No 3 702 77.65 6.27 亲水Hydrophilic 细胞核Nucleus
NPGR2 2224 3 无No 无No 7 737 82.44 7.58 亲水Hydrophilic 细胞核Nucleus

图3

结球甘蓝NPG1、NPGR1、NPGR2基因扩增及序列分析 M: Trans2K plus II DNA marker; A: 结球甘蓝NPG1、NPGR1、NPGR2基因cDNA扩增; B: 结球甘蓝NPG1、NPGR1、NPGR2基因gDNA扩增; C: 结球甘蓝和拟南芥NPG1、NPGR1、NPGR2蛋白TPRs结构域分析; D: 结球甘蓝和拟南芥NPG1-TPR1结构域序列对比。"

图4

结球甘蓝NPG1、NPGR1、NPGR2蛋白进化树分析"

图5

转重组质粒在SD/-Leu/-Trp/X-α-gal/AbA培养基的酵母自激活检测 P: Y187 (AD-T)×Y2H (BK-53); N: Y187 (ADT7-T)×Y2H (BK-Lam); 1: Y2H (BK-CML13)×Y187 (AD-T); 2: Y2H (BK-CML15)× Y187 (AD-T); 3: Y2H (BK-CML20)×Y187 (AD-T); 4: Y2H (BK-CML21)×Y187 (AD-T); 5: Y2H (BK-CML24)×Y187 (AD-T); 6: Y2H (BK-CML25)×Y187 (AD-T); 7: Y2H (BK-CML39)×Y187 (AD-T); 8: Y2H (BK-CML41)×Y187 (AD-T); 9: Y2H (BK-CML42)×Y187 (AD-T); 10: Y2H (BK-CML48) ×Y187 (AD-T); 11: Y2H (BK-CML49)×Y187 (AD-T); 12:Y2H (BK-CML50)×Y187 (AD-T); 13: Y187 (AD-NPG1)×Y2H (BK-53); 14: Y187 (AD-NPGR1)×Y2H (BK-53); 15: Y187 (AD-NPGR2)×Y2H (BK-53)."

表4

结球甘蓝CMLs与NPG1、NPGR1及NPGR2互作分析"

AD\BK CML13 CML15 CML20 CML21 CML24 CML25 CML39 CML41 CML42 CML48 CML49 CML50
NPG1 — + + — — + + + — + + —
NPGR1 — + + — — + + + — + + +
NPGR2 — + + — — + + + — + + +

图6

结球甘蓝CMLs与NPG1、NPGR1、NPGR2互作分析 P: 阳性对照; N: 阴性对照。"

表5

结球甘蓝CMLs与NPG1、NPGR1及NPGR2聚合强度分析"

AD\BK β-半乳糖酶活性 β-galactosidase activity (Miller units)
CML15 CML20 CML25 CML39 CML41 CML48 CML49 CML50
NPG1 5.37 5.85 5.61 5.71 7.77 8.31 6.68 —
NPGR1 4.58 2.31 4.16 3.97 5.31 7.17 2.69 3.65
NPGR2 4.50 4.96 4.82 4.51 6.58 7.70 4.47 5.90

图7

结球甘蓝CMLs(A)和NPG1、NPGR1及NPGR2(B)不同授粉时间表达分析 柱上不同小写字母表示差异显著(P < 0.05)。"

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