Chilling stress, a common abiotic factor, adversely affects the growth and biomass of rice seedlings during the early stages, ultimately reducing the yield. Effective strategies to mitigate these negative impacts are essential for improving rice productivity. The application of nanotechnology in agriculture, particularly nanoparticles (NPs), has shown a promising effect in alleviating chilling stress in plants. This study evaluates the effects of various nanoparticles, ZnO (0, 50, 100, and 200 mg/L), Fe₂O₃ (0, 50, 75, and 100 mg/L), TiO₂ (0, 50, 75, and 100 mg/L), and CeO₂ (0, 50, 75, and 100 mg/L) on the chilling resistance with one control (a water spray) under a normal temperature. Four rice cultivars: LLY-7108 and XZX-6 (Low-temperature-tolerant), and LLY-32 and ZJZ-17 (Low-temperature-susceptible) were tested in this experiment. Rice seedlings were subjected to low temperature conditions (12 h light 14 °C/12 h dark, at 10 °C) for five days, followed by seven days of recovery. The results of this study demonstrate that NPs significantly enhanced seedling height fresh/dry weight and root length compared to untreated controls under chilling stress. NP treatment also reduced the reactive oxygen species (ROS), malondialdehyde (MDA), and proline content, while enhancing superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) activities, thereby mitigating oxidative damage. The four rice varieties exhibited clear signs of rapid growth recovery and positive physiological changes due to NPs’ application. Among the tested cultivars, LLY-7108 showed the most substantial recovery and physiological responses, while ZJZ-17 exhibited the least. The findings of this study indicate that the foliar application of ZnO (100 mg/L), Fe₂O₃ (50 mg/L), TiO₂ (50 mg/L), and CeO₂ (75 mg/L) NPs effectively mitigates chilling stress in rice seedlings, likely by enhancing the antioxidant enzymatic activity while reducing the oxidative damage. This study highlights the potential of NPs as effective agents in reducing the adverse effects of chilling stress on rice.

Cold stress (CS) represents a major environmental factor that adversely affects plant growth, development, and productivity. To cope with low-temperature conditions, plants have evolved sophisticated mechanisms for CS perception and response, mediated through complex cellular signaling networks and physiological processes. Central to these adaptive responses are phytohormones, which function either independently or through synergistic and antagonistic interactions to fine-tune CS tolerance. This review synthesizes current knowledge on the roles of major classical phytohormones and signaling metabolites in regulating CS tolerance in plants. We first outline the molecular mechanisms involved in CS sensing and signal transduction, highlighting the roles of membrane-associated sensors, calcium signaling, and downstream transcriptional networks. Then, we discuss the contributions of key classical phytohormones, including auxin, abscisic acid, ethylene, salicylic acid, cytokinin, jasmonic acid, brassinosteroids, gibberellic acid, strigolactones, and signaling metabolites, including melatonin and gamma-aminobutyric acid, to CS tolerance, highlighting their individual and interacting roles in modulating gene expression regulation, antioxidant defense and physiological adaptations. We also discuss the crosstalk between these hormones, emphasizing the dynamic and often context-dependent nature of their interactions in response to CS. Furthermore, the review highlights recent advances in CRISPR/Cas9-based genome editing strategies targeting phytohormone biosynthesis, signaling, and response pathways to improve CS tolerance in plants. By integrating hormonal signaling, molecular regulation, and modern biotechnological tools, this review provides a comprehensive framework for understanding phytohormone-mediated CS adaptation and offers perspectives for developing climate-resilient crops through genetic and agronomic approaches.

Cold stress (CS) severely limits the growth and productivity of rice (Oryza sativa L.), particularly in temperate regions where abrupt temperature declines frequently occur during early developmental stages. In recent years, nanoparticle (NP) application has emerged as a promising approach for alleviating CS; however, systematic comparisons of different NPs across multiple growth stages remain unclear. This study evaluated the effectiveness and physiological mechanisms of four NPs (Fe₂O₃, ZnO, TiO₂, and CeO₂) in enhancing CS tolerance of rice seedlings using four cultivars with contrasting cold tolerance: two conventional cultivars (ZJZ-17, cold-sensitive; XZX-6, cold-tolerant) and two hybrid cultivars (LLY-7108, cold-tolerant; LLY-32, cold-sensitive). Seedlings were subjected to CS (14 °C day/10 °C night) for 5 days at three developmental stages (14, 21, and 28 days after emergence), followed by a 7-day recovery period under optimal conditions. CS markedly reduced plant height (34.8%), fresh weight (57.2%), dry weight (50.0%), and chlorophyll a and b contents (48%) following recovery. Foliar application of NPs significantly mitigated the adverse effects of CS, with Fe₂O₃ and ZnO showing the highest effectiveness. Fe₂O₃ treatment increased plant height, fresh weight, and dry weight by 25.6%, 43.5%, and 40.6%, respectively, relative to cold-stressed plants, while chlorophyll a and b contents increased by 41.6% and 42.2%. NPs application alleviated oxidative damage by reducing reactive oxygen species (up to 67.4%), malondialdehyde (up to 51.2%), and proline accumulation (up to 60.4%). Enhanced antioxidant defense was evidenced by increased activities of superoxide dismutase (66.6%), peroxidase (59.6%), and catalase (34.3%) under Fe₂O₃ treatment. Yield-related traits also showed significant recovery, with Fe₂O₃ increasing tiller number, spikelets per panicle, and grain yield per plant. The hybrid cultivar LLY-7108 consistently exhibited greater CS tolerance than conventional cultivars, while the cold-sensitive cultivar ZJZ-17 showed the greatest susceptibility. CS imposed at later growth stages (28-day-old seedlings) caused less damage and allowed greater recovery than early-stage stress (14-day-old seedlings). Overall, NP-mediated enhancement of photosynthesis and antioxidant capacity significantly improves CS tolerance and yield performance in rice, with Fe₂O₃ NPs emerging as a promising strategy for mitigating CS. These findings provide practical insights for rice cultivation in regions prone to chilling events and contribute to the development of nanoparticle-based approaches for rice production under climate stress.

Rice faces increasing threats from climate change, driven by heat, salinity, and drought stresses. Although researchers have extensively studied individual stresses, growing evidence indicates that combined stresses elicit unique physiological, biochemical, and molecular responses that single-stress studies cannot predict. This review comprehensively synthesizes current knowledge of the integrated signaling networks and adaptive mechanisms underlying rice responses to heat, salinity, and drought stress, with particular emphasis on stress crosstalk and multi-stress responses. We examine the roles of mechanosensitive ion channels, calcium signaling, salt overly sensitive (SOS) pathway, heat shock response regulators, mitogen-activated protein kinase (MAPK) cascades, and ABA-dependent and -independent signaling networks in stress tolerance systems. Furthermore, we discuss major adaptive responses involving reactive oxygen species (ROS) homeostasis, osmoprotectant accumulation, antioxidant defense systems, and phytohormone crosstalk involving abscisic acid, auxin, salicylic acid, cytokinin, brassinosteroids, ethylene, and gibberellins. Particular attention is given to the detrimental effects of combined stresses on growth, development, and yield. In addition, recent advances in enhancing stress tolerance through CRISPR/Cas9-mediated genome editing, marker-assisted breeding, and molecular marker technologies are also highlighted. Despite significant progress, critical knowledge gaps remain regarding primary stress-sensing mechanisms, epigenetic stress memory, organelle-specific signaling, and the translation of laboratory findings to field conditions. By integrating current knowledge across molecular, physiological, and breeding perspectives, we provide a comprehensive integrated signaling network model and crosstalk among signaling pathways for understanding rice adaptation to combined stresses and highlight potential strategies for enhancing crop resilience and global food security.

Cold stress significantly impairs the rice (Oryza sativa L.) growth and yield, particularly in temperate regions where abrupt temperature fluctuations ofte