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作物学报 ›› 2026, Vol. 52 ›› Issue (3): 764-779.doi: 10.3724/SP.J.1006.2026.53047

所属专题： 玉米：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米ZmPFK基因家族全基因组鉴定及响应胁迫表达分析

孟成(), 王哲()   

  1. 作物抗逆与高效生产全国重点实验室 / 农业农村部西北旱区玉米生物学与遗传育种重点实验室 / 西北农林科技大学农学院, 陕西杨凌 712100
  • 收稿日期:2025-07-09 接受日期:2025-11-18 出版日期:2026-03-12 网络出版日期:2025-12-03
  • 通讯作者: *王哲, E-mail: wangzhe8636@nwafu.edu.cn
  • 作者简介:E-mail: mengc@nwafu.edu.cn
  • 基金资助:
    国家自然科学基金项目(32101699);中央高校科研业务费项目(Z1090221024)

Genome-wide identification and expression analysis of the ZmPFK gene family under biotic and abiotic stresses in maize

Meng Cheng(), Wang Zhe()   

  1. State Key Laboratory of Crop Stress Resistance and High-Efficiency Production / Key Laboratory of Maize Biology and Genetic Breeding in Arid Area of Northwest Region, Ministry of Agriculture and Rural Affairs / College of Agronomy, Northwest A&F University, Yangling 712100, Shaanxi, China
  • Received:2025-07-09 Accepted:2025-11-18 Published:2026-03-12 Published online:2025-12-03
  • Contact: *王哲, E-mail: wangzhe8636@nwafu.edu.cn
  • Supported by:
    National Natural Science Foundation of China(32101699);Fundamental Research Funds for the Central Universities(Z1090221024)

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摘要：

磷酸果糖激酶(phosphofructokinase, PFK)是糖酵解途径中的关键调控酶, 作为该代谢过程的限速酶, 它主导着细胞内葡萄糖代谢的速率。与此同时, 己糖激酶不仅能够作为糖受体启动下游糖信号传导, 还可整合多种外界信号, 进而激活其他信号转导途径, 参与调节植物对外界环境变化的响应机制。系统分析玉米PFK家族成员在基因组上的分布、结构、进化以及表达特性, 对于深入研究该家族的生物学功能具有重要意义。本研究基于玉米B73 (RefGen_v4)基因组信息以及转录组数据等, 通过生物信息学手段对玉米B73自交系磷酸果糖激酶(PFK)家族成员进行全基因组鉴定并对其理化性质、系统发育关系、基因结构、启动子顺式作用元件以及表达特性进行了分析。结果表明, 玉米B73自交系共有20个PFK家族成员, 它们主要分为ATP依赖性的磷酸果糖激酶(phosphofructokinase, PFK)和焦磷酸盐依赖性的磷酸果糖激酶(phosphotransferase, PFP), 且不均匀的分布于各个染色体上。共线性分析表明, 玉米PFK基因与单子叶植物水稻(Oryza sativa)具有多对同源基因, 而与双子叶植物拟南芥(Arabidopsis thaliana)不存在直系同源基因。启动子顺式作用元件预测表明, PFK基因具有一些与生长发育和逆境响应的结合元件。转录组数据以及qRT-PCR表明, 各成员在组织表达与胁迫处理下存在不同的表达水平, 并且发现了多个响应非生物胁迫(盐胁迫、干旱胁迫、热胁迫和冷胁迫)与生物胁迫(玉蜀黍平脐蠕孢菌、玉蜀黍凸脐蠕孢菌和禾谷镰孢胁迫)的ZmPFK基因, 表明该家族成员在响应逆境胁迫中具有重要作用。将重组质粒ZmPFK10-pET32a转入大肠杆菌BL21中, 诱导表达后的重组菌株ZmPFK10-pET32a在700 mmol L-1 NaCl和45℃高温条件下的生长情况优于对照菌株。蛋白互作网络预测与富集分析发现, 与ZmPFK蛋白存在互作的3个核心蛋白均参与糖酵解活动。上述结果为PFK家族成员的生物学功能研究提供了理论基础。

关键词: 玉米, PFK, 基因家族, 糖酵解, 表达分析

Abstract:

Phosphofructokinase (PFK) is a key regulatory enzyme in the glycolytic pathway, functioning as the rate-limiting enzyme that controls the pace of intracellular glucose metabolism. In parallel, hexokinase not only serves as a sugar sensor initiating downstream sugar signaling, but also integrates various external cues, thereby activating other signaling pathways and contributing to the regulation of plant responses to environmental stimuli. A systematic investigation of the gene structure, evolutionary relationships, and expression patterns of PFK family members in maize is crucial for advancing our understanding of their biological functions. In this study, a genome-wide identification of PFK family members was conducted in the maize inbred line B73 using bioinformatics approaches. The physicochemical properties, phylogenetic relationships, gene structures, cis-acting regulatory elements, and expression profiles were comprehensively analyzed. Twenty PFK genes were identified in B73, categorized into ATP-dependent phosphofructokinases (PFKs) and pyrophosphate-dependent phosphotransferases (PFPs), and were found to be unevenly distributed across different chromosomes. Collinearity analysis revealed multiple homologous genes between maize and rice (Oryza sativa), but none with Arabidopsis thaliana. Promoter analysis indicated the presence of cis-regulatory elements associated with growth, development, and stress responses. Transcriptome data and qRT-PCR analysis demonstrated differential expression of ZmPFK genes across tissues and under various stress treatments. Several ZmPFK genes responded to abiotic stresses (salt, drought, heat, and cold) as well as biotic stresses caused by Cochliobolus heterostrophus, Exserohilum turcicum, and Fusarium graminearum, suggesting a potential role for the PFK family in stress adaptation. Moreover, the recombinant plasmid ZmPFK10-pET32a was introduced into Escherichia coli BL21, and the resulting strain exhibited enhanced growth under 700 mmol L-1 NaCl and 45℃, compared to the control. Protein interaction network prediction and enrichment analysis further revealed that three core proteins interacting with ZmPFK are involved in glycolysis. These findings provide a theoretical foundation for future functional characterization of PFK family members in maize.

Key words: maize, PFK, gene family, glycolysis, expression analysis

表1

ZmPFK基因qRT-PCR引物信息"

基因名称Gene name 引物名称Primer name 引物序列Primer sequence (5'-3')
ZmPFK5 ZmPFK5-F CCGAACCCCCTCCAAGACAA
ZmPFK5-R GCTGGTCTTGTGAACAACGATCT
ZmPFK6 ZmPFK6-F GGGGAGGGTGGACTATTTGA
ZmPFK6-R GCTTGGAATAGCACGGATCA
ZmPFK10 ZmPFK10-F CAAGATCGGCGTTGTGCTCTC
ZmPFK10-R AGCTGGGCCTCCCTTAAATCC
ZmPFK12 ZmPFK12-F CCAGCCTATTCAACTGTCAAG
ZmPFK12-R CCACACACCAGTTCCCTGATAA
ZmPFK19 ZmPFK19-F TGGTTGAGACCGAAATGGAC
ZmPFK19-R CCCAGGGTTTCGGTAGATGT
ZmEF-1α ZmEF-1α-F TGGGCCTACTGGTCTTACTACTGA
ZmEF-1α-R ACATACCCACGCTTCAGATCCT

表2

ZmPFK10基因引物信息"

基因名称Gene name 引物名称Primer name 引物序列Primer sequence (5'-3')
ZmPFK10 ZmPFK10-F gccatggctgatatcggatccATGGCGGCGGCGGCGGTG
ZmPFK10-R acggagctcgaattcggatccTAGCGCAGCACCGAGTTCC

表3

玉米PFK基因家族基本信息"

基因名称
Gene name		 基因ID
Gene ID		 染色体位置
Chromosome localization		 氨基酸长度
Amino acid length (aa)		 分子量
Molecular weight (D)		 等电点
Isoelectric point		 不稳定指数
Instability index		 类群
Group
ZmPFK1 Zm00001d032172_T004 Chr.1 527 58,192.72 8.23 37.93 PFK
ZmPFK2 Zm00001d032386_T003 Chr.1 618 67,282.41 6.79 43.40 PFP
ZmPFK3 Zm00001d033002_T001 Chr.1 525 57,475.48 7.04 30.57 PFK
ZmPFK4 Zm00001d005756_T002 Chr.2 567 62,602.72 8.16 36.79 PFK
ZmPFK5 Zm00001d039499_T002 Chr.3 528 57,293.35 7.94 33.10 PFK
ZmPFK6 Zm00001d043563_T002 Chr.3 764 83,089.25 6.00 42.66 PFK
ZmPFK7 Zm00001d051629_T003 Chr.4 624 67,945.26 7.24 40.27 PFP
ZmPFK8 Zm00001d017830_T001 Chr.5 618 67,473.40 7.85 38.79 PFP
ZmPFK9 Zm00001d036206_T004 Chr.6 552 60,828.26 8.22 36.37 PFK
ZmPFK10 Zm00001d037278_T001 Chr.6 564 61,051.75 6.30 32.83 PFP
ZmPFK11 Zm00001d037910_T001 Chr.6 526 57,323.34 7.64 35.38 PFK
ZmPFK12 Zm00001d038775_T001 Chr.6 575 63,221.28 6.00 40.19 PFK
ZmPFK13 Zm00001d020006_T001 Chr.7 544 59,642.11 5.69 47.32 PFP
ZmPFK14 Zm00001d020866_T001 Chr.7 147 15,528.91 9.10 21.54 PFK
ZmPFK15 Zm00001d008816_T002 Chr.8 531 57,650.68 7.92 34.33 PFK
ZmPFK16 Zm00001d010375_T001 Chr.8 538 58,039.24 7.60 33.38 PFK
ZmPFK17 Zm00001d044754_T001 Chr.9 580 62,806.72 5.96 31.91 PFP
ZmPFK18 Zm00001d045326_T004 Chr.9 542 59,506.65 8.22 35.55 PFK
ZmPFK19 Zm00001d045919_T001 Chr.9 476 52,076.02 6.64 40.21 PFP
ZmPFK20 Zm00001d025659_T001 Chr.10 477 51,231.13 6.21 34.54 PFK

图1

ZmPFKs蛋白亚细胞定位 PFK蛋白亚细胞定位结果, 数值越高表明定位概率越大。cysk: 细胞骨架; cyto: 细胞质; chlo: 叶绿体; nucl: 细胞核; mito: 线粒体; vacu: 液泡; plas: 细胞膜; extr: 细胞外基质。"

图2

玉米、拟南芥和水稻中PFK进化关系 A: PFK系统发育树, AtPFK (拟南芥), OsPFK (水稻), ZmPFK (玉米)。红、黄、蓝、绿、紫分别表示I~V五种类群; B:PFK蛋白序列比对, 热图中颜色越深表明相似度越高。"

图3

玉米PFK家族保守基序和基因结构分析 A: 保守基序; B: 基因结构。"

图4

玉米PFK基因在染色体上的分布"

图5

拟南芥、水稻和玉米的共线性分析与玉米PFK家族共线性分析 A: 玉米与拟南芥基因组间的共线性关系; B: 玉米与水稻基因组间的共线性关系; C: 玉米基因组内共线性关系。灰线代表物种间基因组所有共线性关系, 红线表示PFK基因在物种间的共线性关系。Zm: 玉米; At: 拟南芥; Os: 水稻。渐变色注图表示染色体上每10 kb基因数量。"

图6

玉米PFK基因启动子顺式作用元件预测 W-box: WRKY转录因子特异结合位点; ABRE: 脱落酸响应元件; TGA-element: 生长素响应元件; CCAAT-box: 以核心序列5'-CCAAT- 3'命名; MBS、MBSI、MYB: MYB转录因子结合位点; STRE: 应激响应元件; MYC: MYC转录因子特异结合位点; WRE3: 伤口响应元件; TCA: 以元件中包含TCA序列命名; CGTCA-motif: 以核心序列5'-CGTCA-3'命名; LTR: 低温响应元件; P-box: 赤霉素响应元件; GARE-motif: 赤霉素响应元件; AuxRR-core: 生长素响应核心元件; TATC-box: 以核心序列5'-TATC-3'命名; TC-rich repeats: 以富含TC重复序列命名; CARE: C和G框相关联的顺式作用元件。"

图7

玉米PFK基因组织特异性表达"

图8

ZmPFK基因在非生物胁迫与生物胁迫下的表达 A: 非生物胁迫, Salt为盐胁迫, Heat为热胁迫, Drought为干旱胁迫, Cold为冷胁迫; B: 生物胁迫, C.h为玉蜀黍平脐蠕孢菌, E.t为玉蜀黍凸脐蠕孢菌, F.g为禾谷镰孢。"

图9

ZmPFK基因实时荧光定量结果 C. h: 玉蜀黍平脐蠕孢菌侵染。t检验进行显著性分析。ns: 差异不显著。"

图10

ZmPFK10的扩增 第1泳道为2000 bp的Marker; 第2泳道为加入模板的克隆; 第3泳道为未加模板的阴性对照。"

图11

ZmPFK10在大肠杆菌BL21中的功能验证"

图12

ZmPFK蛋白互作网络和核心互作蛋白的KEGG富集分析"

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