Genome Wide Identification, Molecular Characterization, and Gene Expression Analyses of Grapevine NHX Antiporters Suggest Their Involvement in Growth, Ripening, Seed Dormancy, and Stress Response

  • Mariem Ayadi
  • Viviana Martins
  • Rayda Ben Ayed
  • Rania Jbir
  • Molka Feki
  • Rim Mzid
  • Hernâni Géros
  • Sami Aifa
  • Mohsen HananaEmail author
Original Article


Plant NHX antiporters are critical for cellular pH, Na+, and K+ homeostasis and salt tolerance. Even though their genomic and functional studies have been conducted in many species, the grapevine NHX family has not been described yet. Our work highlights the presence of six VvNHX genes whose phylogenetic analysis revealed their classification in two distinct groups: group I vacuolar (VvNHX1–5) and group II endosomal (VvNHX6). Several cis-acting regulatory elements related to tissue-specific expression, transcription factor binding, abiotic/biotic stresses response, and light regulation elements were identified in their promoter. Expression profile analyses of VvNHX genes showed variable transcription within organs and tissues with diverse patterns according to biochemical, environmental, and biotic treatments. All VvNHXs are involved in berry growth, except VvNHX5 that seems to be rather implicated in seed maturation. VvNHX4 would be more involved in floral development, while VvNHX2 and 3 display redundant roles. QPCR expression analyses of VvNHX1 showed its induction by NaCl and KNO3 treatments, whereas VvNHX6 was induced by ABA application and strongly repressed by PEG treatment. VvNHX1 plays a crucial role in a bunch of grape developmental steps and adaptation responses through mechanisms of phyto-hormonal signaling. Overall, VvNHX family members could be valuable candidate genes for grapevine improvement.


NHX antiporter Berry Grapevine Salinity Ripening Dormancy Vacuolar Endosomal 



This research work was financially supported by the Ministry of Higher Education and Scientific Research of Tunisia. We are grateful to Mr Riadh BEN MARZOUG for his helps with some software.

Author Contributions

AM, HM, and HG designed the study. AM, BAR, JR, FM, LV, AS, MR, and HM performed data analyses. AM, MV, and HG performed the experiments. AM, MV, and HM analyzed the data and wrote the manuscript. All authors read and approved the final manuscript.

Compliance with Ethical Standards

Conflict of interest

The authors declare no conflicts of interest.

Supplementary material

10528_2019_9930_MOESM1_ESM.docx (15 kb)
Supplementary material 1 (DOCX 26 kb)
10528_2019_9930_MOESM2_ESM.xlsx (20 kb)
Supplementary material 2 (xlsx) (XLSX 21 kb)
10528_2019_9930_MOESM3_ESM.xlsx (15 kb)
Supplementary material 3 Online Resource 3: Cq values generated by QPCR analysis (XLSX 15 kb)
10528_2019_9930_MOESM4_ESM.pptx (111 kb)
Supplementary material 4 Online Resource 4: Melting curves s generated by QPCR analysis (PPTX 111 kb)


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Authors and Affiliations

  • Mariem Ayadi
    • 1
    • 2
  • Viviana Martins
    • 3
    • 4
    • 5
  • Rayda Ben Ayed
    • 2
  • Rania Jbir
    • 6
  • Molka Feki
    • 2
  • Rim Mzid
    • 1
  • Hernâni Géros
    • 3
    • 4
    • 5
  • Sami Aifa
    • 2
  • Mohsen Hanana
    • 1
    Email author
  1. 1.Laboratory of Extremophile Plants, Center of Biotechnology of Borj-CédriaHammam-lifTunisia
  2. 2.Laboratory of Molecular and Cellular Screening Processes, Center of Biotechnology of SfaxUniversity of SfaxSfaxTunisia
  3. 3.Centro de Biologia Molecular E Ambiental (CBMA), Departamento de Biologia, Escola de CiênciasUniversidade Do MinhoBragaPortugal
  4. 4.Centro de Investigação E de Tecnologias Agro-Ambientais E Biológicas (CITAB)Universidade de Trás-Os-Montes E Alto DouroVila RealPortugal
  5. 5.Centro de Engenharia Biológica (CEB)Universidade Do MinhoBragaPortugal
  6. 6.Laboratory of Plant Biotechnologies Applied To Crop ImprovementNational School of Engineering of Sfax (ENIS)SfaxTunisia

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