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Novel application of CKX inhibitors and chlorpromazine overcomes recalcitrance in spinach tissue culture

Ashkan Hodaei (UGent) and Stefaan Werbrouck (UGent)
(2025) PLANT CELL TISSUE AND ORGAN CULTURE. 161(3).
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Abstract
Spinach (Spinacia oleracea), a highly valuable leafy vegetable, presents significant challenges for in vitro regeneration due to its recalcitrant nature, hindering genetic improvement efforts. This study aimed to develop an efficient regeneration protocol for spinach by investigating the effects of novel small molecules in tissue culture media. In addition to traditional plant growth regulators, we incorporated a cytokinin oxidase inhibitor (3TFM-2HE) and chlorpromazine (CPZ), a combination that has not been previously reported for plant tissue culture. Various combinations and concentrations of these regulators were evaluated for direct and indirect organogenesis. While direct organogenesis proved unsuccessful, a two-step protocol involving callus induction followed by shoot regeneration yielded positive results. To initiate the process, competent callus formation was induced through a combination of 10 mu M Naphthaleneacetic Acid (NAA), 10 mu M 2-isopentenyladenosine (2iPR), 1 mu M 3TFM-2HE, and 50 mu M CPZ. Subsequently, the introduction of 5 mu M Gibberellic Acid (GA3) stimulated successful shoot regeneration. This is the first in vitro regeneration report for the commercially important spinach cultivar Palco, with no prior tissue culture studies. This protocol provides a valuable tool for spinach improvement and demonstrates the potential of cytokinin oxidase inhibitors and CPZ for enhancing regeneration in other recalcitrant species.
Keywords
Callus induction, Shoot induction, Regeneration, 2iPR, NAA, Competent callus, Palco spinach, PLANT-REGENERATION, CYTOKININ HOMEOSTASIS, HYDROGEN-PEROXIDE, CALMODULIN ANTAGONISTS, GIBBERELLIC-ACID, AUXIN, ROOT, EXPRESSION, CALLUS, RICE

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Citation

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MLA
Hodaei, Ashkan, and Stefaan Werbrouck. “Novel Application of CKX Inhibitors and Chlorpromazine Overcomes Recalcitrance in Spinach Tissue Culture.” PLANT CELL TISSUE AND ORGAN CULTURE, vol. 161, no. 3, 2025, doi:10.1007/s11240-025-03098-z.
APA
Hodaei, A., & Werbrouck, S. (2025). Novel application of CKX inhibitors and chlorpromazine overcomes recalcitrance in spinach tissue culture. PLANT CELL TISSUE AND ORGAN CULTURE, 161(3). https://doi.org/10.1007/s11240-025-03098-z
Chicago author-date
Hodaei, Ashkan, and Stefaan Werbrouck. 2025. “Novel Application of CKX Inhibitors and Chlorpromazine Overcomes Recalcitrance in Spinach Tissue Culture.” PLANT CELL TISSUE AND ORGAN CULTURE 161 (3). https://doi.org/10.1007/s11240-025-03098-z.
Chicago author-date (all authors)
Hodaei, Ashkan, and Stefaan Werbrouck. 2025. “Novel Application of CKX Inhibitors and Chlorpromazine Overcomes Recalcitrance in Spinach Tissue Culture.” PLANT CELL TISSUE AND ORGAN CULTURE 161 (3). doi:10.1007/s11240-025-03098-z.
Vancouver
1.
Hodaei A, Werbrouck S. Novel application of CKX inhibitors and chlorpromazine overcomes recalcitrance in spinach tissue culture. PLANT CELL TISSUE AND ORGAN CULTURE. 2025;161(3).
IEEE
[1]
A. Hodaei and S. Werbrouck, “Novel application of CKX inhibitors and chlorpromazine overcomes recalcitrance in spinach tissue culture,” PLANT CELL TISSUE AND ORGAN CULTURE, vol. 161, no. 3, 2025.
@article{01JW747B1VXYPZ8J8RMPTAW5TQ,
  abstract     = {{Spinach (Spinacia oleracea), a highly valuable leafy vegetable, presents significant challenges for in vitro regeneration due to its recalcitrant nature, hindering genetic improvement efforts. This study aimed to develop an efficient regeneration protocol for spinach by investigating the effects of novel small molecules in tissue culture media. In addition to traditional plant growth regulators, we incorporated a cytokinin oxidase inhibitor (3TFM-2HE) and chlorpromazine (CPZ), a combination that has not been previously reported for plant tissue culture. Various combinations and concentrations of these regulators were evaluated for direct and indirect organogenesis. While direct organogenesis proved unsuccessful, a two-step protocol involving callus induction followed by shoot regeneration yielded positive results. To initiate the process, competent callus formation was induced through a combination of 10 mu M Naphthaleneacetic Acid (NAA), 10 mu M 2-isopentenyladenosine (2iPR), 1 mu M 3TFM-2HE, and 50 mu M CPZ. Subsequently, the introduction of 5 mu M Gibberellic Acid (GA3) stimulated successful shoot regeneration. This is the first in vitro regeneration report for the commercially important spinach cultivar Palco, with no prior tissue culture studies. This protocol provides a valuable tool for spinach improvement and demonstrates the potential of cytokinin oxidase inhibitors and CPZ for enhancing regeneration in other recalcitrant species.}},
  articleno    = {{73}},
  author       = {{Hodaei, Ashkan and Werbrouck, Stefaan}},
  issn         = {{0167-6857}},
  journal      = {{PLANT CELL TISSUE AND ORGAN CULTURE}},
  keywords     = {{Callus induction,Shoot induction,Regeneration,2iPR,NAA,Competent callus,Palco spinach,PLANT-REGENERATION,CYTOKININ HOMEOSTASIS,HYDROGEN-PEROXIDE,CALMODULIN ANTAGONISTS,GIBBERELLIC-ACID,AUXIN,ROOT,EXPRESSION,CALLUS,RICE}},
  language     = {{eng}},
  number       = {{3}},
  pages        = {{12}},
  title        = {{Novel application of CKX inhibitors and chlorpromazine overcomes recalcitrance in spinach tissue culture}},
  url          = {{http://doi.org/10.1007/s11240-025-03098-z}},
  volume       = {{161}},
  year         = {{2025}},
}
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