Abstract
Light quality is a critical determinant in controlled environment agriculture, yet information regarding the interactive effects of spectral composition and grafting on tomato seedling performance remains limited. Moving beyond the assumption that rootstock vigor is solely a static trait, we hypothesized that above-ground spectral cues significantly modulate rootstock efficiency. We investigated the synergistic effects of grafting (Solanum lycopersicum ‘Maxifort’) and various LED spectra (monochromatic Red, Blue, White, and Red:Blue 70:30) on morphological architecture, photosynthetic potential, and the stoichiometric balance of mineral nutrients in tomato seedlings. Our results reveal a critical interaction: while grafting alone alleviated specific nutritional deficits (N, K, Mg) under suboptimal monochromatic red light, the rootstock’s capacity to maximize the uptake of key elements—particularly phosphorus and calcium—was fully realized only under the synergistic Red:Blue spectrum. This study provides empirical evidence that integrating the R70:B30 spectrum with grafting not only improves growth but also optimizes resource acquisition. These findings offer a novel approach to optimizing transplant quality and establish a robust protocol for producing resilient transplants in modern nurseries. Future research should focus on unraveling the molecular pathways underlying this light-rootstock communication.
Similar content being viewed by others
Data availability
The data presented in this study are available on request from the corresponding author.
References
Echeverría-Pérez, E. G. et al. Recent developments of nanomaterials in crop growth and production: The case of the tomato (Solanum lycopersicum). Agronomy 15, 1716 (2025).
Govindasamy, R., Ceylan, R. F. & Özkan, B. Global tomato production: price sensitivity and policy impact in Mexico, Türkiye, and the United States. Horticulturae 11, 84 (2025).
Shafe, M. O., Gumede, N. M., Nyakudya, T. T. & Chivandi, E. Lycopene: A potent antioxidant with multiple health benefits. J. Nutr. Metab. 2024, (2024).
Balali, A., Fathzadeh, K., Askari, G. & Sadeghi, O. Dietary intake of tomato and lycopene, blood levels of lycopene, and risk of total and specific cancers in adults: A systematic review and dose–response meta-analysis of prospective cohort studies. Front. Nutr. 12, (2025).
Gallegos-Cedillo, V. M. et al. Analysis of global research on vegetable seedlings and transplants and their impacts on product quality. J. Sci. Food Agric. 104, 4950–4965 (2024).
Khopade, R. Y. et al. Vegetable grafting: a scientific innovation to enhance productivity and profitability of tomato growers under climate change. Front. Agron. 7, (2025).
Lee, J. M. et al. Current status of vegetable grafting: Diffusion, grafting techniques, automation. Sci. Hortic. (Amsterdam) 127, 93–105 (2010).
Awazade, A. S. & Verma, D. Advancing vegetable grafting: A comprehensive review of techniques, challenges, and the future of automated solutions. J. Sci. Res. Reports 30, 517–530 (2024).
Sahoo, B. et al. The art of grafting: Elevating vegetable production to new height. E-Planet 22, 24–39 (2024).
Colla, G., Suãrez, C. M. C., Cardarelli, M. & Rouphael, Y. Improving nitrogen use efficiency in melon by grafting. HortScience 45, 559–565 (2010).
Rouphael, Y., Cardarelli, M., Colla, G. & Rea, E. Yield, mineral composition, water relations, and water use efficiency of grafted mini-watermelon plants under deficit irrigation. HortScience 43, 730–736 (2008).
Venema, J. H., Dijk, B. E., Bax, J. M., van Hasselt, P. R. & Elzenga, J. T. M. Grafting tomato (Solanum lycopersicum) onto the rootstock of a high-altitude accession of Solanum habrochaites improves suboptimal-temperature tolerance. Environ. Exp. Bot. 63, 359–367 (2008).
Gruda, N. S., Samuolienė, G., Dong, J. & Li, X. Environmental conditions and nutritional quality of vegetables in protected cultivation. Compr. Rev. Food Sci. Food Saf. 24, (2025).
Han, R., Lin, R., Zhou, Y. & Thomas, H. R. Here comes the sun: integration of light, temperature, and auxin during herbaceous plant grafting. Planta 261, 124 (2025).
Wu, W. et al. The role of light in regulating plant growth, development and sugar metabolism: a review. Front. Plant Sci. 15, (2024).
Sena, S., Kumari, S., Kumar, V. & Husen, A. Light emitting diode (LED) lights for the improvement of plant performance and production: A comprehensive review. Curr. Res. Biotechnol. 7, 100184 (2024).
Gruda, N. S. et al. Advancing protected cultivation: A pathway for nutrient-rich vegetables. CRC. Crit. Rev. Plant Sci. 44, 88–116 (2025).
Azizi, S. et al. Photobiology, photosynthesis, and plant responses under artificial lighting in controlled environment agriculture. Sci. Hortic. 349, 114248 (2025).
Heuvelink, E. et al. Tomato in the spotlight: Light regulation of whole-plant physiology in tomato. J. Exp. Bot. https://doi.org/10.1093/jxb/eraf315 (2025).
Meiramkulova, K. et al. The efficiency of led irradiation for cultivating high-quality tomato seedlings. Sustain. 13, 9426 (2021).
Zhang, Y. et al. Manipulation of artificial light environment improves plant biomass and fruit nutritional quality in tomato. J. Adv. Res. 75, 79–93 (2024).
Kim, D. et al. Effects of Supplemental LED Lighting based on Solar Irradiation and Carbon Dioxide Enrichment on Photosynthesis, Growth, and Yield of Greenhouse-Grown Tomato Plants. J. People, Plants, Environ. 28, 1–12 (2025).
Li, B. et al. The impact of light intensities on the phenotypic parameters of cucumber seedlings at three developmental stages. Agronomy 14, 627 (2024).
Aliniaeifard, S. et al. Light in controlled environment agriculture. Int. J. Veg. Sci. 31, 615–621 (2025).
Bueno, P. M. C. & Vendrame, W. A. Wavelength and light intensity affect macro- and micronutrient uptake, stomata number, and plant morphology of common bean (Phaseolus vulgaris L.). Plants 13, 441 (2024).
Wang, T. et al. Effects of red and blue light on the growth, photosynthesis, and subsequent growth under fluctuating light of cucumber seedlings. Plants 13, 1668 (2024).
Arif, A. Bin et al. Application of red and blue LED light on cultivation and postharvest of tomatoes (Solanum lycopersicum L.). Scientifica (Cairo). 2024, (2024).
He, W. et al. Response of blue light in different proportions on the growth & flowering in sunflower. Sci. Hortic. 338, 113689 (2024).
Trivellini, A., Toscano, S., Romano, D. & Ferrante, A. The role of blue and red light in the orchestration of secondary metabolites, nutrient transport and plant quality. Plants 12, 2026 (2023).
Wang, J., Lu, W., Tong, Y. & Yang, Q. Leaf morphology, photosynthetic performance, chlorophyll fluorescence, stomatal development of lettuce (Lactuca sativa L.) exposed to different ratios of red light to blue light. Front. Plant Sci. 7, (2016).
Fukuda, N., Ajima, C., Yukawa, T. & Olsen, J. E. Antagonistic action of blue and red light on shoot elongation in petunia depends on gibberellin, but the effects on flowering are not generally linked to gibberellin. Environ. Exp. Bot. 121, 102–111 (2016).
Hogewoning, S. W. et al. Blue light dose-responses of leaf photosynthesis, morphology, and chemical composition of Cucumis sativus grown under different combinations of red and blue light. J. Exp. Bot. 61, 3107–3117 (2010).
Naznin, M. T., Lefsrud, M., Gravel, V. & Azad, M. O. K. Blue light added with red LEDs enhance growth characteristics, pigments content, and antioxidant capacity in lettuce, Spinach, Kale, Basil, and sweet pepper in a controlled environment. Plants 8, 93 (2019).
Gallegos-Cedillo, V. M. et al. An in-depth analysis of sustainable practices in vegetable seedlings nurseries: A review. Sci. Hortic. 334, 113342 (2024).
Glowacka, B. The effect of blue light on the height and habit of the tomato (Lycopersicon esculentum Mill.) transplant. Folia Hortic. 16, 3–10 (2004).
Kaiser, E. et al. Adding blue to red supplemental light increases biomass and yield of greenhouse-grown tomatoes, but only to an optimum. Front. Plant Sci. 9, (2019).
Roosta, H. R., Samadi, A. & Bikdeloo, M. Different cultivation systems and foliar application of calcium nanoparticles affect the growth and physiological characteristics of pennyroyal (Mentha pulegium L.). Sci. Rep. 13, 20334 (2023).
Yousef, A. F. et al. Effects of light spectrum on morphophysiological traits of grafted tomato seedlings. PLoS ONE 16, e0250210 (2021).
Bantis, F., Koukounaras, A., Siomos, A. S., Fotelli, M. N. & Kintzonidis, D. Bichromatic red and blue LEDs during healing enhance the vegetative growth and quality of grafted watermelon seedlings. Sci. Hortic. (Amsterdam). 261, 109000 (2020).
Ke, X., Yoshida, H., Hikosaka, S. & Goto, E. Effect of red and blue light versus white light on fruit biomass radiation-use efficiency in dwarf tomatoes. Front. Plant Sci. 15, (2024).
Nguyen, D. T. P., Kitayama, M., Lu, N. & Takagaki, M. Improving secondary metabolite accumulation, mineral content, and growth of coriander (Coriandrum sativum L.) by regulating light quality in a plant factory. J. Hortic. Sci. Biotechnol. 95, 356–363 (2020).
Seif, M. et al. Monochromatic red light during plant growth decreases the size and improves the functionality of stomata in chrysanthemum. Funct. Plant Biol. 48, 515–528 (2021).
Cao, H. et al. Genome-wide identification and expression analysis of the cryptochromes reveal the CsCRY1 role under low-light-stress in cucumber. Front. Plant Sci. 15, (2024).
Dierck, R., Dhooghe, E., Van Huylenbroeck, J., Van Der Straeten, D. & De Keyser, E. Light quality regulates plant architecture in different genotypes of Chrysanthemum morifolium Ramat. Sci. Hortic. (Amsterdam) 218, 177–186 (2017).
Fang, L. et al. Plant growth and photosynthetic characteristics of soybean seedlings under different LED lighting quality conditions. J. Plant Growth Regul. 40, 668–678 (2021).
Yang, F. O., Mao, J. F., Wang, J., Zhang, S. & Li, Y. Transcriptome analysis reveals that red and blue light regulate growth and phytohormone metabolism in Norway spruce [Picea abies (L.) Karst.]. PLoS ONE 10, e0127896 (2015).
Yu, X., Liu, H., Klejnot, J. & Lin, C. The cryptochrome blue light receptors. Arab. B. 8, e0135 (2010).
Kayal, W. E. et al. Molecular events of apical bud formation in white spruce, Picea glauca. Plant Cell Environ. 34, 480–500 (2011).
Kurepin, L. V. et al. Interactions between plant hormones and light quality signaling in regulating the shoot growth of arabidopsis thaliana seedlings. Botany 90, 237–246 (2012).
Lau, O. S. & Deng, X. W. Plant hormone signaling lightens up: Integrators of light and hormones. Curr. Opin. Plant Biol. 13, 571–577 (2010).
Liu, J. & van Iersel, M. W. Photosynthetic physiology of blue, green, and red light: Light intensity effects and underlying mechanisms. Front. Plant Sci. https://doi.org/10.3389/fpls.2021.619987 (2021).
Moosavi-Nezhad, M. et al. Blue light improves photosynthetic performance during healing and acclimatization of grafted watermelon seedlings. Int. J. Mol. Sci. 22, 8043 (2021).
Si, C. et al. Effects of LED light quality combinations on growth and leaf colour of tissue culture-generated plantlets in Sedum rubrotinctum. Hortic. Sci. Technol. 42, 53–67 (2024).
Paradiso, R. & Proietti, S. Light-quality manipulation to control plant growth and Photomorphogenesis in greenhouse horticulture: The state of the art and the opportunities of modern LED systems. J. Plant Growth Regul. 41, 742–780 (2022).
Kumar, P., Rouphael, Y., Cardarelli, M. & Colla, G. Effect of nickel and grafting combination on yield, fruit quality, antioxidative enzyme activities, lipid peroxidation, and mineral composition of tomato. J. Plant Nutr. Soil Sci. 178, 848–860 (2015).
Malekzadeh Shamsabad, M. R. et al. Supplemental light application can improve the growth and development of strawberry plants under salinity and alkalinity stress conditions. Sci. Rep. 12, 9272 (2022).
An, H. et al. Effects of nutrition and light quality on the growth of southern highbush blueberry (Vaccinium corymbosum L.) in an advanced plant factory with artificial lighting (PFAL). Horticulturae 9, 287 (2023).
Brazaitytė, A. et al. Effect of blue light percentage on mineral elements content in Brassica microgreens. Acta Hortic. 1271, 119–125 (2020).
Amoozgar, A., Mohammadi, A. & Sabzalian, M. R. Impact of light-emitting diode irradiation on photosynthesis, phytochemical composition and mineral element content of lettuce cv. Grizzly. Photosynthetica 55, 85–95 (2017).
Savvas, D. et al. Effects of three commercial rootstocks on mineral nutrition, fruit yield, and quality of salinized tomato. J. Plant Nutr. Soil Sci. 174, 154–162 (2011).
Singh, H. et al. Grafting tomato as a tool to improve salt tolerance. Agronomy 10, 263 (2020).
Suh, D. H. et al. Characterization of metabolic changes under low mineral supply (N, K, or Mg) and supplemental LED lighting (red, blue, or red⇓blue combination) in perilla frutescens using a metabolomics approach. Molecules 25, 4714 (2020).
Brazaitytė, A. et al. Effect of different ratios of blue and red led light on brassicaceae microgreens under a controlled environment. Plants 10, 801 (2021).
Breive, K. et al. High-power light-emitting diode based facility for plant cultivation. J. Phys. D. Appl. Phys. 38, 3182–3187 (2005).
Wang, S. et al. The red/blue light ratios from light-emitting diodes affect growth and flower quality of Hippeastrum hybridum ‘Red Lion’. Front. Plant Sci. 13, (2022).
Yan, J. et al. Light quality regulates plant biomass and fruit quality through a photoreceptor-dependent HY5-LHC/CYCB module in tomato. Hortic. Res. 10, (2023).
Sager, J. C., Smith, W. O., Edwards, J. L. & Cyr, K. L. Photosynthetic efficiency and phytochrome photoequilibria determination using spectral data. Trans. Am. Soc. Agric. Eng. 31, 1882–1889 (1988).
Tahmasebi, F. Physiological investigation of the irrigation effect with salty water from NaCl and CaCl2 sources on three genotype of Canola (Brassica napus L.) in Ahvaz climate. (2010).
John Ryan, G. E. and A. R. Soil and plant analysis laboratory manual. Sustain. 11, (2001).
Hoel, B. O. & Solhaug, K. A. Effect of irradiance on chlorophyll estimation with the Minolta SPAD-502 leaf chlorophyll meter. Ann. Bot. 82, 389–392 (1998).
Funding
Open Access funding enabled and organized by Projekt DEAL.
Author information
Authors and Affiliations
Contributions
Conceptualization, S.S. and R.S.; methodology, H.A. and R.S.; formal analysis, S.S.; investigation, S.S.; data curation, S.S.; writing—original draft preparation, S.S.; writing—review and editing, H.A., R.S., and NSG; supervision, H.A. and R.S.; project administration, R.S. and NSG. All authors have read and agreed to the published version of the manuscript.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Informed consent
Not applicable.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
About this article
Cite this article
Soltani, S., Aroiee, H., Salehi, R. et al. The synergistic effect of grafting and LED light quality on enhancing the mineral nutrition and growth performance of tomato seedlings. Sci Rep (2026). https://doi.org/10.1038/s41598-026-38960-3
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-026-38960-3
