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作物学报 ›› 2025, Vol. 51 ›› Issue (12): 3224-3237.doi: 10.3724/SP.J.1006.2025.54063

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

马铃薯CEP基因家族鉴定及StCEP2小肽调控马铃薯不定根生长和耐盐性的功能分析

张慧敏(), 党亚茹, 桑佳男, 焦文静, 孙春怡, 庄佳牧, 魏玉树, 张超, 田鹏, 刘柏林, 宋银()   

  1. 西北农林科技大学农学院 / 作物抗逆与高效生产全国重点实验室, 陕西杨凌 712100
  • 收稿日期:2025-05-19 接受日期:2025-09-10 出版日期:2025-12-12 网络出版日期:2025-09-25
  • 通讯作者: *宋银, E-mail: yin.song@nwafu.edu.cn
  • 作者简介:E-mail: zhanghuimin2022@nwafu.edu.cn
  • 基金资助:
    本研究由国家重点研发计划项目(2022YFF1002700);陕西省重点研发计划项目(2021LLRH-07);陕西省重点研发计划项目(2024NC-YBXM-002);大学生创新创业训练计划项目(S202410712556)

Genome-wide identification of CEP gene family in potato (Solanum tuberosum L.) and functional analysis of StCEP2 in regulating adventitious root growth and salt tolerance

ZHANG Hui-Min(), DANG Ya-Ru, SANG Jia-Nan, JIAO Wen-Jing, SUN Chun-Yi, ZHUANG Jia-Mu, WEI Yu-Shu, ZHANG Chao, TIAN Peng, LIU Bai-Lin, SONG Yin()   

  1. College of Agronomy, Northwest A&F University / State key Laboratory for Crop Stress Resistance and High-efficiency Production, Yangling 712100, Shaanxi, China
  • Received:2025-05-19 Accepted:2025-09-10 Published:2025-12-12 Published online:2025-09-25
  • Contact: *E-mail: yin.song@nwafu.edu.cn
  • Supported by:
    National Key R&D Program of China(2022YFF1002700);Key R&D Program in Shaanxi Province(2021LLRH-07);Key R&D Program in Shaanxi Province(2024NC-YBXM-002);Undergraduate Training Program for Innovation and Entrepreneurship(S202410712556)

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摘要： CEP (C-terminally encoded peptide)编码一类在C-末端附近具有保守基序的分泌小肽, 属于多成员的基因家族, 在调控植物根系生长、氮素营养及逆境胁迫中起重要作用。本研究对马铃薯(Solanum tuberosum) CEP基因(StCEP)家族成员进行了鉴定, 并对StCEP家族成员的理化特征、保守基序、系统进化关系和启动子顺式作用元件进行了分析; 利用RNA-seq数据分析了StCEP家族成员在马铃薯不同组织以及响应不同环境信号的表达模式, 并筛选受盐胁迫显著诱导的StCEP基因; 最后利用外源施加合成StCEP小肽的方法对StCEP小肽的耐盐性功能进行了验证。结果表明, 在马铃薯基因组中共鉴定到11个StCEP家族成员, 分布在4条不同的染色体上, 其编码的蛋白含有1~9个数目不等的CEP基序; StCEP家族成员与拟南芥、番茄CEP成员相互嵌合形成2个大的分支, 各分支分别包含21个和26个CEP成员; 马铃薯与拟南芥之间仅存在1对CEP基因共线性关系, 而与亲缘关系较近的番茄之间存在12对CEP基因共线性关系; StCEP基因的启动子区域主要包括光响应、激素响应、植物生长发育响应和逆境胁迫响应的顺式作用元件; StCEP基因的表达具有明显的组织特异性且受氮素、激素(BAP、IAA、GA3和ABA)、不同生物(致病疫霉、BABA和BTH)与非生物(NaCl)逆境因子的诱导; 外施StCEP2小肽可以促进马铃薯不定根的生长并增强马铃薯的耐盐性。本研究为后续深入解析StCEP基因家族成员在马铃薯生长发育与响应环境信号过程中的功能提供了参考信息。

关键词: 马铃薯, CEP小肽, 外施小肽, 不定根生长, 耐盐性

Abstract:

C-terminally encoded peptide (CEP) genes encode a class of secreted small peptides characterized by conserved structural domains near the C-terminus. These genes form a multi-member family that plays crucial roles in regulating various physiological processes in plants, including root development, nitrogen uptake, and responses to environmental stresses. In this study, members of the CEP gene family in potato (Solanum tuberosum, StCEP) were identified and analyzed for their physicochemical properties, conserved motifs, phylogenetic relationships, and promoter cis-acting elements. Expression patterns of StCEP genes were examined across different potato tissues and under various environmental conditions using RNA-seq data. Additionally, the function of salt-induced StCEP genes was validated through exogenous application of synthetic StCEP peptides. A total of 11 StCEP genes were identified in the potato genome, distributed across four chromosomes. The encoded StCEP proteins contained between 1 and 9 CEP motifs, with variability in motif number among members. Phylogenetic analysis revealed that StCEP family members clustered with CEP genes from Arabidopsis thaliana and Solanum lycopersicum, forming two major clades comprising 21 and 26 members, respectively. Synteny analysis showed one syntenic gene pair between potato and A. thaliana, and 12 syntenic pairs between potato and S. lycopersicum. Promoter analysis indicated that StCEP genes are primarily regulated by light-responsive, hormone-responsive, developmental, and stress-related cis-elements. Expression profiling revealed that StCEP genes exhibit tissue-specific expression and are responsive to nitrogen availability, phytohormones (BAP, IAA, GA3, and ABA), and both biotic (e.g., Phytophthora infestans, BABA, BTH) and abiotic (e.g., NaCl) stress factors. Furthermore, exogenous application of the StCEP2 peptide promoted adventitious root formation and enhanced salt tolerance in potato plantlets under in vitro conditions. This study offers valuable insights into the functional roles of StCEP genes in regulating plant development and responses to environmental stress, laying a foundation for future functional genomics and crop improvement research.

Key words: potato, CEP peptide, exogenous peptide assay, adventitious root growth, salt-stress tolerance

图1

StCEP基因的染色体分布及基因结构 A: StCEP基因在染色体上的分布; B: StCEP基因的基因结构。"

表1

StCEP基因家族成员的理化特征"

基因名称
Gene name		 基因编号
Gene ID		 氨基酸
数目
No. of amino acids		 分子量
Molecular weight (kD)		 等电点
pI		 不稳定指数Instability index 脂肪指数Aliphatic index 亚细胞定位
Subcellular
localization		 信号肽
Signal peptide
StCEP1 Soltu.DM.02G008180.1 90 9.77 9.43 34.71 85.67 胞外Extracellular +
StCEP2 Soltu.DM.02G028150.1 91 9.96 9.60 34.85 90.00 线粒体Mitochondrial +
StCEP3 Soltu.DM.02G029100.1 108 11.60 9.90 47.87 95.65 胞外Extracellular +
StCEP4 Soltu.DM.02G029110.1 143 15.44 8.79 50.91 74.97 胞外Extracellular +
StCEP5 Soltu.DM.03G005620.1 97 10.31 6.90 42.52 98.45 胞外Extracellular +
StCEP6 Soltu.DM.03G005950.1 123 13.61 7.02 33.15 100.49 胞外Extracellular +
StCEP7 Soltu.DM.03G005980.1 119 13.28 9.05 54.43 71.18 细胞核Nuclear +
StCEP8 Soltu.DM.07G023480.1 354 33.90 9.80 30.04 49.10 细胞核Nuclear +
StCEP9 Soltu.DM.07G023490.1 211 20.96 7.95 33.68 55.88 细胞核Nuclear -
StCEP10 Soltu.DM.07G023500.1 240 25.51 9.46 50.68 69.67 细胞核, 细胞质, 线粒体
Nuclear, cytoplasmic,
mitochondrial		 -
StCEP11 Soltu.DM.08G014640.1 80 9.17 9.60 29.46 52.25 细胞核, 细胞质
Nuclear, cytoplasmic		 -

图2

StCEP蛋白中CEP基序的鉴定与分析 A: StCEP蛋白中CEP基序的比对分析。B: 拟南芥、番茄和马铃薯CEP蛋白的CEP基序的Weblogo图。C: StCEP蛋白N端信号肽的剪切位点; 红色箭头所指为预测的信号肽剪切位点。D: CEP基序的系统发育树; CEP基序被划分为2组, 即Group I和Group II。E: Group I和Group II内CEP基序的Weblogo图。"

图3

马铃薯、番茄和拟南芥CEP家族成员的系统发育树与共线性关系 A: 马铃薯、番茄和拟南芥CEP家族成员的系统发育树; B: StCEP基因分别与拟南芥和番茄CEP基因之间的共线性关系。具有共线性关系的CEP家族成员用相同颜色显示。"

图4

StCEP基因家族成员启动子区的顺式作用元件 方框中的数字表示每个StCEP基因启动子区具有顺式作用元件的个数。"

图5

StCEP基因在马铃薯不同组织的表达模式 热图方框中的数值为基因表达量指标TPM (transcripts per kilobase million)进行log2(TPM+1)标准化处理所得的值。"

图6

StCEP基因在氮素、激素、生物逆境因子和盐胁迫下的表达模式 A: 在低氮(0.2 mmol L-1)或高氮(4 mmol L-1)处理下, StCEP基因在匍匐茎、地上部分和根中的表达模式; B: 在植物激素10 μmol L-1 BAP、10 μmol L-1 IAA、50 μmol L-1 GA3和50 μmol L-1 ABA处理24 h后StCEP基因的表达模式; C: 在生物逆境因子致病疫霉(P. infestans)侵染、化学诱导剂β-氨基丁酸和苯并噻二唑处理后StCEP基因在马铃薯叶片中的表达情况; D: 在200 mmol L-1 NaCl处理0、3、6、12和24 h后StCEP基因的表达模式。"

图7

外施StCEP2小肽促进马铃薯不定根生长和增强马铃薯对盐胁迫的耐受性 A: 外施1 μmol L-1 StCEP2p小肽培养15 d后, 马铃薯不定根生长以及在盐胁迫(150 mmol L-1)条件下马铃薯不定根的表型; 标尺代表 3 cm。B: 外施1 μmol L-1 StCEP2p小肽培养15 d后, 马铃薯不定根表面积以及在盐胁迫(150 mmol L-1)条件下马铃薯不定根表面积。C: 外施1 μmol L-1 StCEP2p小肽培养15 d后, 试管苗的鲜重以及在盐胁迫(150 mmol L-1)条件下试管苗的鲜重。数值代表试管苗鲜重或不定根表面积的平均值±SEM。不同字母代表不同处理样本间的试管苗鲜重或不定根表面积具有显著差异性(P < 0.05)。"

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