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Article

Scion–Rootstock Interactions Enhance Freezing Stress Resilience in Citrus reticulata Through Integrated Antioxidant Defense and Carbon–Nitrogen Metabolic Adjustments

by
Alaiha Asif
1,†,
Shahid Iqbal
1,†,
Carlos Eduardo Aucique-Perez
1,
KeAndre Leaks
1,
Rashad Mukhtar Balal
2,
Matthew Mattia
3,
John M. Chater
4 and
Muhammad Adnan Shahid
1,*
1
Horticultural Science Department, North Florida Research and Education Center, IFAS/University of Florida, Quincy, FL 32351, USA
2
Department of Horticulture, College of Agriculture, University of Sargodha, Sargodha 40100, Pakistan
3
United States Horticultural Research Laboratory, United States Department of Agriculture, Agricultural Research Service, 2001 South Rock Road, Fort Pierce, FL 34945, USA
4
Horticultural Science Department, Citrus Research and Education Center, IFAS/University of Florida, Lake Alfred, FL 33850, USA
*
Author to whom correspondence should be addressed.
These authors contributed equally to this work.
Plants 2025, 14(19), 3029; https://doi.org/10.3390/plants14193029
Submission received: 29 August 2025 / Revised: 22 September 2025 / Accepted: 24 September 2025 / Published: 30 September 2025
(This article belongs to the Special Issue Rootstock Influence on Crop Regulation)

Abstract

Frequent and increasingly severe freezing events threaten citrus production in northern Florida, underscoring the need for strategies that enhance freezing resilience in citrus cultivars. Grafting scions onto tolerant rootstocks provides a physiologically integrative approach to improve stress tolerance. This study aims to elucidate how these interactions modulate physiological and metabolic responses under freezing stress, thereby identifying mechanisms that contribute to enhanced freeze resilience in citrus. Here, we grafted Citrus reticulata (cv. UF-950) onto eight rootstocks (Bitters, Blue-1, C-146, Sour Orange, UFR07TC, UFR09TC, UFR5, and US942) to evaluate scion–rootstock interactions under normal (20 °C) and freezing (−6 °C) conditions. Freezing stress caused a sharp increase in oxidative stress markers, lipid peroxidation, and membrane damage while reducing photosynthetic performance across most combinations. Antioxidant capacity, osmolyte accumulation, and carbon–nitrogen metabolic responses varied significantly among rootstocks, revealing strong genotype-dependent modulation of scion physiology. Among the tested combinations, UF-950 grafted onto UFR5 displayed the highest freezing tolerance, characterized by robust activation of antioxidant enzymes, elevated proline and glycine betaine accumulation, reduced oxidative damage, and sustained carbon–nitrogen metabolic fluxes under freezing stress. These results demonstrate that rootstock genotype governs the extent of scion defense activation and metabolic homeostasis under freezing conditions. Our findings identify UFR5 as a promising rootstock for enhancing freezing resilience in citrus and provide mechanistic insight into how scion–rootstock interaction orchestrates integrative stress tolerance pathways. Future work should focus on multi-omics dissection of rootstock-mediated signaling networks and long-term field validation to optimize rootstock selection for enhanced cold resilience under variable climatic conditions.
Keywords: antioxidants; freeze tolerance; photosynthesis; rootstock; reactive oxygen species antioxidants; freeze tolerance; photosynthesis; rootstock; reactive oxygen species

Share and Cite

MDPI and ACS Style

Asif, A.; Iqbal, S.; Aucique-Perez, C.E.; Leaks, K.; Balal, R.M.; Mattia, M.; Chater, J.M.; Shahid, M.A. Scion–Rootstock Interactions Enhance Freezing Stress Resilience in Citrus reticulata Through Integrated Antioxidant Defense and Carbon–Nitrogen Metabolic Adjustments. Plants 2025, 14, 3029. https://doi.org/10.3390/plants14193029

AMA Style

Asif A, Iqbal S, Aucique-Perez CE, Leaks K, Balal RM, Mattia M, Chater JM, Shahid MA. Scion–Rootstock Interactions Enhance Freezing Stress Resilience in Citrus reticulata Through Integrated Antioxidant Defense and Carbon–Nitrogen Metabolic Adjustments. Plants. 2025; 14(19):3029. https://doi.org/10.3390/plants14193029

Chicago/Turabian Style

Asif, Alaiha, Shahid Iqbal, Carlos Eduardo Aucique-Perez, KeAndre Leaks, Rashad Mukhtar Balal, Matthew Mattia, John M. Chater, and Muhammad Adnan Shahid. 2025. "Scion–Rootstock Interactions Enhance Freezing Stress Resilience in Citrus reticulata Through Integrated Antioxidant Defense and Carbon–Nitrogen Metabolic Adjustments" Plants 14, no. 19: 3029. https://doi.org/10.3390/plants14193029

APA Style

Asif, A., Iqbal, S., Aucique-Perez, C. E., Leaks, K., Balal, R. M., Mattia, M., Chater, J. M., & Shahid, M. A. (2025). Scion–Rootstock Interactions Enhance Freezing Stress Resilience in Citrus reticulata Through Integrated Antioxidant Defense and Carbon–Nitrogen Metabolic Adjustments. Plants, 14(19), 3029. https://doi.org/10.3390/plants14193029

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