Plant responses to heat stress and advances in mitigation strategies

Abstract

High-temperature stress is a major abiotic constraint limiting plant growth and agricultural productivity. While its adverse effects are well documented, most studies have examined individual species or isolated physiological mechanisms. This review provides a comprehensive comparative analysis of heat stress responses across four major crops - barley (Hordeum vulgare), rice (Oryza sativa), maize (Zea mays), and tomato (Solanum lycopersicum), alongside the model plant Arabidopsis thaliana, focusing on their morphological, physiological, and biochemical adaptations as well as current mitigation strategies. Morphological assessments reveal that root traits are more heat-sensitive than shoot length, biomass, or germination rate. Physiologically, all species exhibit reduced photosynthetic rate and PSII efficiency (Fv/Fm), though stomatal conductance and transpiration responses vary. Biochemically, the accumulation of reactive oxygen species (ROS) and antioxidant activity exhibit species- and stress-dependent regulation, with both upregulation and downregulation observed. Among mitigation approaches, seed priming emerges as a cost-effective strategy, while miRNA-mediated regulation shows strong potential for developing heat-tolerant cultivars. This synthesis highlights critical knowledge gaps and outlines future directions for enhancing crop resilience in the face of rising temperatures.

1 Introduction

Recent urbanization and population growth have placed unprecedented pressure on agricultural systems, while climate change has emerged as one of the critical threats to global crop productivity (Kousar et al., 2021; Benitez-Alfonso et al., 2023; Farooq et al., 2023; Terán et al., 2024). The accelerating pace of global warming is particularly alarming, with 2024 projected to be the warmest year on record, approximately 1.55°C above pre-industrial levels (WMO, 2024). Climate models suggest 66% probability that at least one year between 2024 and 2028 will temporarily exceed the 1.5°C threshold, with sustained exceedance likely by the early 2030s (IPCC, 2023). This warming trend poses severe challenges to agriculture, where heat stress, exacerbated by the increasing frequency of extreme weather events, significantly compromises crop yields and quality (Mirón et al., 2023; Janni et al., 2024). Meta-analyses indicate that each 1°C increase in temperature reduces the yield of major crops by 3–7% (Challinor et al., 2014; Zhao et al., 2017; Tran et al., 2025), creating substantial risks to global food security as demand is projected to rise by 50% by 2050 (FAO, 2021). Addressing these challenges requires both innovative mitigation strategies and a deeper understanding of plant responses to heat stress to facilitate breeding of climate-resilient varieties (Kousar et al., 2021; Terán et al., 2024).

Although plant responses to heat stress have been extensively studied, most reviews focus on individual species or specific mechanisms, limiting broad interspecific comparisons (Wu et al., 2019; Jacott and Boden, 2020; Medina et al., 2021; Elakhdar et al., 2022; El-Sappah et al., 2022; LI et al., 2022; Singh et al., 2022; Ren et al., 2023; Sarma et al., 2023; Ruan et al., 2024). Similarly, comprehensive studies on Oryza sativa (rice) emphasize varietal differences in thermotolerance, hormonal regulation, and physiological adaptations (Wu et al., 2019; Ren et al., 2023; Sarma et al., 2023). In Zea mays (maize), heat stress responses have been dissected from molecular, transcriptional, and agronomic perspectives, while studies on Solanum lycopersicum (Tomato) focus on reactive oxygen species (ROS) signaling and biochemical adjustments (Medina et al., 2021; El-Sappah et al., 2022; LI et al., 2022; Singh et al., 2022; Ruan et al., 2024). Despite these advances, a systematic comparison across species is lacking. Our review addresses this critical knowledge gap through a comprehensive interspecific analysis of five representative species – Hordeum vulgare, Oryza sativa, Zea mays, Solanum lycopersicum, and Arabidopsis thaliana – spanning both monocots and dicots. These species were selected based on their agronomic importance, diverse growth habits, photosynthetic pathways, ecological adaptations, and availability of heat stress response data, with A. thaliana included as a well-established model plant. We systematically evaluate their morphological, physiological, and biochemical adaptation to heat stress across different experimental systems.

Beyond assessing stress responses, this review critically compares current mitigation strategies, which are often examined in isolation rather than across multiple species (Liu et al., 2022; Zhang et al., 2022; Feng et al., 2023; Iqbal et al., 2023; Louis et al., 2023; Ma and Hu, 2023). Seed priming, for example, enhances thermotolerance by boosting antioxidant activity, inducing stress memory, and even conferring transgenerational resilience (Liu et al., 2022; Louis et al., 2023). Chemical and nutritional treatments mitigate heat stress through osmotic regulation and antioxidant defense activation, while microbial biostimulants improve tolerance via beneficial plant–microbe interactions (Feng et al., 2023; Iqbal et al., 2023). Additionally, miRNAs have gained attention as key post-transcriptional regulators of heat stress signaling and adaptation (Zhang et al., 2022; Ma and Hu, 2023). However, most of these studies evaluate these strategies in a species-specific context, obscuring broader trends. Here, we systematically analyze four major mitigation approaches across five species, assessing comparative efficacy and providing a unified perspective on heat stress management in crops.

2 Morphological changes in plants under heat stress

Each plant species possesses unique optimal temperature conditions, indispensable for sustaining normal growth and development (Krutovsky et al., 2025). These conditions can exhibit considerable variation depending on the species’ genetic background and ecological origins. When the temperatures exceed this range, morphological disruptions occur, including reduced seed germination, abnormal root and shoot development, and overall growth impairment. These changes serve as critical indicators of heat stress damage, particularly in economically important crops such as H. vulgare, O. sativa, Z. mays, S. lycopersicum, and the model plant Arabidopsis thaliana (Ito et al., 2009; Hussain et al., 2019a; Naz et al., 2022).

2.1 Seed germination response to high temperatures

Seed germination is one of the most heat-sensitive stages of plant development, making it a key marker for assessing heat stress effects (Scandalios et al., 2000; Mei and Song, 2010; Liu et al., 2015). Supra-optimal temperatures disrupt germination, essential for successful plant establishment and growth (Table 1) (Baskin and Baskin, 2000; Brändel, 2004). Above 30°C, germination rates decline sharply, and under extreme heat (>45°C), germination may be severely reduced or completely inhibited (Liu et al., 2015).

As shown in Table 1, the extent of germination reduction varies among plant species. For example, germination rates decrease by 95.3% in barley cv. Pijiu, 90% in rice cv. Peiai, 50% in tomato cv. C38, and 75% in Arabidopsis cv. Columbia (Labouriau and Osborn, 1984; Mei and Song, 2010; Liu et al., 2015; Afshar et al., 2016; Tokić et al., 2023). In contrast, heat-tolerant maize cultivars, w64a, r6-67, and dn-6, exhibit a more moderate decline (~40%), highlighting genotypic variation in thermotolerance (Scandalios et al., 2000).

However, laboratory-based germination studies may not fully reflect field conditions, where interacting environmental factors (temperature fluctuations, humidity, light, soil nutrients) influence seed behavior (Klupczyńska and Pawłowski, 2021; Qaderi, 2023). While controlled experiments provide valuable insights into specific physiological mechanisms, they often fail to capture ecological complexity and may overestimate seed germination resilience. Field conditions present a more challenging environment where seeds encounter unpredictable stress combinations – including diurnal temperature fluctuations, variable soil moisture, and biotic interactions – that collectively alter germination outcomes compared to uniform laboratory conditions. Additionally, germination is only significantly affected under extreme heat, showing low sensitivity at moderate stress levels. Thus, relying solely on germination data may be insufficient for predicting overall plant productivity under heat stress, and it requires a broader focus, incorporating subsequent morphological parameters.

2.2 Impact of heat stress on root and shoot growth

Elevated temperatures severely suppress root and shoot expansion, leading to reduced biomass accumulation. Impaired root function under heat stress disrupts water and nutrient uptake, further exacerbating growth limitations (Koevoets et al., 2016; Taratima et al., 2022). Among the analyzed species, root fresh weight (RFW) emerges as one of the most heat-sensitive root parameters (Table 2). Arabidopsis, rice, and barley show severe reductions in RFW, ranging from 52% to 70% under thermal stress, while maize and tomato exhibit more moderate declines of 29% and 39%, respectively (Abo Gamar et al., 2019; Hussain et al., 2019a; Guo et al., 2022; Naz et al., 2022; Baloch et al., 2024). This pattern extends to reduce root dry weight (RDW), where similar percentage reductions suggest parallel impacts on both biomass accumulation and water retention capacity (Xia et al., 2021; Guo et al., 2022; Macabuhay et al., 2022; Naz et al., 2022; Baloch et al., 2024).

While both root and shoot biomass are temperature-sensitive, shoot fresh weight (SFW) generally displays greater stress resilience than root parameters (Table 3). Significant SFW reductions (29-50%) occur in barley, tomato, Arabidopsis, and rice, contrasting with maize’s notably higher tolerance (only 3.2-8% reduction) (Hussain et al., 2016; Abo Gamar et al., 2019; Zhou et al., 2019; Naz et al., 2022; Baloch et al., 2024). This suggests that despite the negative effects of heat stress, some plant species exhibit a higher capacity to maintain shoot and leaf biomass production. This interspecific variation in shoot response is further evidenced by shoot dry weight (SDW) measurements, where Arabidopsis and barley suffer substantial losses (20-25%), compared to more modest impacts on maize (3%), tomato (8%), and rice (11%), highlighting fundamental differences in thermotolerance mechanisms among species (Ito et al., 2009; Hussain et al., 2016; Zhou et al., 2019; Macabuhay et al., 2022; Naz et al., 2022).

In addition to reducing biomass, heat stress profoundly influences root and shoot elongation (Table 4). The extent of these reductions varies among species, reflecting differences in their ability to tolerate high temperatures. Root length reductions vary from 12% in rice cv. Vandana, 26% in barley, 48% in tomato, to 55% in Arabidopsis cv. Columbia (Sailaja et al., 2014; Yang et al., 2017; Guo et al., 2022; Naz et al., 2022). A similar trend is observed for shoot length, with reductions of 5%-10% in Arabidopsis, 14% in barley and rice, and 59.1% in tomato under heat-stress conditions (Sailaja et al., 2014; Ibañez et al., 2017; Guo et al., 2022; Naz et al., 2022). Maize cv. SD609 again demonstrates exceptional thermal stability, with only 35% root length reduction and minimal shoot length impact (6-8%), reinforcing its status as a heat-resilient crop (Hussain et al., 2016; Xia et al., 2021). This relative stability in shoot growth suggests species-specific adaptations to thermal stress.

These morphological alterations directly impair critical physiological functions, including nutrient uptake, water balance, and overall metabolic homeostasis, collectively compromising normal growth and development (Yamaura et al., 2021; Barten et al., 2022; Pérez-Bueno et al., 2022; Visakorpi et al., 2023). The synergistic effects of diminished biomass production, restricted elongation growth, and impaired hydration status ultimately lead to severe yield penalties, particularly in heat-sensitive species (Figure 1). At the cellular level, these macroscopic changes reflect underlying disruptions in hormone signaling, oxidative homeostasis, and carbon metabolism, with suppressed photosynthetic capacity (manifested through reduced leaf expansion, limited biomass production, and stunted growth) representing a key factor exacerbating heat stress impacts on plant performance and productivity (Greer and Weedon, 2012; Brunel-Muguet et al., 2015; Mahmood et al., 2021, 2022; Bernacchi et al., 2023; Burroughs et al., 2023; Batool et al., 2025; Li et al., 2025; Secomandi et al., 2025).

Figure 1

3 Formatting physiological changes in plants under heat stress

High temperatures significantly impair critical physiological processes, particularly photosynthetic efficiency, which serves as a key indicator of photosystem functionality. Essential parameters affected include the maximum quantum efficiency of photosystem II (PSII), stomatal conductance, and transpiration rate - all crucial for proper plant growth and development (Rollins et al., 2013; Zheng et al., 2025). These head-induced disruptions lead to diminished carbon assimilation, reduced water use efficiency, and overall growth inhibition.

3.1 Photosynthetic parameters

One of the primary consequences of heat stress is the inactivation of Rubisco, accompanied by reductions in chlorophyll content and PSII efficiency, collectively causing a substantial decline in photosynthetic activity (Wijewardene et al., 2021; Qu et al., 2023; Scafaro et al., 2023; Zahra et al., 2023). Experimental data (Table 5) demonstrate consistent reductions in both photosynthesis rate and PSII maximum quantum yield (Fv/Fm) across species: photosynthesis decreases by 30-50% in barley, 23% in rice, 16.6% in maize, 20% in tomato, and 16% in Arabidopsis (Vile et al., 2012; Zhou et al., 2017; Hussain et al., 2019a; Yang et al., 2020; Mahalingam et al., 2022). Similarly, Fv/Fm values decrease under stress, with reductions of 66.6% in rice, 20% in Arabidopsis, 7.5% in barley, 4% in maize, and 7% in tomato (Rollins et al., 2013; Halter et al., 2017; Zhou et al., 2017; Yang et al., 2018; Zhang et al., 2018; Doğru, 2021). These species-specific response patterns highlight distinct thermotolerance mechanisms among plants, with maize exhibiting relatively greater photosynthetic stability under heat stress compared to more sensitive species like rice and barley.

The detrimental effects of heat stress extend beyond simple reductions in chlorophyll content and PSII efficiency, significantly disrupting critical regulatory mechanisms including stomatal conductance and transpiration rate (Marchin et al., 2023; Zahra et al., 2023; Falcioni et al., 2024; Miao et al., 2025). Under elevated temperature, stomatal conductance – the crucial regulator of gas exchange and water loss – demonstrates a biphasic response: an initial increase to promote transpirational cooling, followed by a decline during prolonged heat exposure (Faralli et al., 2022; Marchin et al., 2022, 2023; Liang et al., 2023). This dynamic pattern exhibits considerable interspecies variation, reflecting distinct evolutionary adaptations to thermal stress. The intricate relationship between stomatal behavior, transpiration efficiency, and photosynthetic performance reveals the sophisticated nature of plant thermoregulation, where maintaining an optimal balance between water conservation and carbon fixation becomes paramount for survival under heat stress conditions.

3.2 Stomatal and transpiration responses to heat stress

Among the key physiological responses to heat stress, stomatal conductance is crucial in regulating gas exchange and transpiration. Studies show this parameter exhibits the most pronounced increase under elevated temperatures, with barley cultivars demonstrating a 50–80% rise after 5–15 days of exposure to 28°C and 38°C (Mahalingam et al., 2022). Similarly, other species also demonstrate an increase to varying degrees: rice (20%), Arabidopsis (11%), maize (10–30%), and decrease tomato (33%) (Table 6) (Jin et al., 2011; Hussain et al., 2016; Zhou et al., 2017; Yang et al., 2020; Mahalingam et al., 2022). These disparities highlight the importance of considering experimental context when interpreting heat stress responses, particularly regarding tissue specificity (leaves vs. whole seedlings), plant developmental stage, duration and intensity of heat exposure, and species-specific tolerance.

While short-term heat exposure typically enhances stomatal conductance, prolonged stress often leads to their decline, as observed in multiple tree species, including various oaks (Quercus macrocarpa Michx, Q. muehlenbergii Engl., Q. stellata), sweetgum (Liquidambar styraciflua), and other species like Vitis vinifera L., Ficus insipida Willd., and Ochroma pyramidale (Hamerlynck and Knapp, 1996; Gunderson et al., 2002; Slot et al., 2016; Zhang et al., 2023; Veloo et al., 2025). Plants employ contrasting strategies to cope with heat stress - some temporarily increase stomatal opening for evaporative cooling while others rapidly close stomata to conserve water (Huang et al., 2025; Luo et al., 2025; Wu et al., 2025). These adaptive responses significantly influence water-use efficiency, photosynthetic performance, and long-term stress resilience.

Plants activate multiple mechanisms to counter prolonged heat stress, including transpirational cooling and stomatal regulation, which help balance water loss and cellular homeostasis (Mathur et al., 2014). However, while stomatal closure prevents dehydration, it simultaneously limits CO₂ uptake, reducing photosynthetic efficiency and carbon assimilation (Hasanuzzaman et al., 2013; Faralli et al., 2022). Species-specific transpiration patterns reflect this balance, with the increase of 35-62% in rice, 30% in barley, 12% in Arabidopsis, 6–9% in maize, and 7% in tomato (Table 6). As highlighted by Jagadish et al., elevated transpiration coupled with effective leaf cooling can significantly mitigate heat stress impacts on plant growth and development (Jagadish et al., 2015). This underscores the critical need to optimize stomatal dynamics in crop breeding programs to enhance heat tolerance and maintain agricultural productivity under increasingly frequent and intense heat stress conditions.

While transpiration provides temporary heat relief, prolonged exposure disrupts fundamental photosynthetic processes (Figure 2). Elevated temperatures trigger complex responses, including altered reactive oxygen species (ROS) dynamics, reduced PSII (Fv/Fm) efficiency, and suppressed photosynthetic activity, collectively limiting carbon assimilation (Batool et al., 2025; Li et al., 2025; Ostendarp et al., 2025). The resulting decline in photosynthetic efficiency directly constrains plant growth and productivity. A comprehensive understanding of the interplay between stomatal behavior, transpiration, and photosynthesis is therefore essential for developing climate-resilient crops in our warming world (Zeng et al., 2017).

Figure 2

4 Biochemical changes in plants under heat stress

Environmental stressors such as extreme temperatures, drought, and salinity frequently induce oxidative stress in plants, leading to cellular damage (Gill and Tuteja, 2010; Li et al., 2023; Nurbekova et al., 2024; Rao and Zheng, 2025; Secomandi et al., 2025). Among these, heat stress poses a particularly severe threat to plant physiology by triggering excessive accumulation of reactive oxygen species (ROS) (Fortunato et al., 2023; Rao and Zheng, 2025). These highly reactive molecules, including hydrogen peroxide (H₂O₂) and superoxide (O₂^-), disrupt cellular processes and contribute to oxidative stress, resulting in membrane damage, impaired metabolism, and overall decline in plant health (Gill and Tuteja, 2010; Samat et al., 2024; Jardim-Messeder et al., 2025; Kračun et al., 2025).

4.1 Reactive oxygen species accumulation under heat stress

Heat stress induces substantial increases in ROS levels across various plant species, contributing to cellular damage and oxidative stress. Studies demonstrate that heat exposure leads to significant rises in H₂O₂ and O₂^- levels, though notable interspecific variation. For instance, H₂O₂ increases by approximately 245% in some species, while barley, maize, and tomato cultivars exhibit a more moderate increase of around 50% (Zhao et al., 2018; Ahammed et al., 2019; Ayub et al., 2021; Zahra et al., 2022). Similarly, O₂^- levels surge by 200% in rice, tomato, and maize cultivars (Table 7) (Zhao et al., 2018; Zhang et al., 2019; Jahan et al., 2022). However, contrasting reports indicate decreases in O₂^- and H₂O₂ by 26% and 27%, respectively, in barley shoots under heat stress (Zhanassova et al., 2021). These discrepancies may stem from differences in experimental conditions, as mentioned previously. Such variations emphasize the complexity of oxidative stress responses and underscore the importance of standardized experimental approaches when comparing interspecific ROS regulation under heat stress, and highlight the need for further investigation into oxidative stress responses.

A major consequence of excessive ROS accumulation is lipid peroxidation, which compromises membrane integrity. Malondialdehyde (MDA), a lipid peroxidation byproduct, serves as a reliable biomarker for oxidative membrane damage. Under heat stress, MDA levels increase by 151% and 150% in rice and tomato cultivars, respectively, with the most pronounced changes observed in rice cultivars (XQZ and G46) during the reproductive stage (Zhao et al., 2018; Ahammed et al., 2019). Comparatively, barley, Arabidopsis, and maize cultivars show increases of 100%, 65%, and 100%, respectively (Table 7), confirming MDA as a consistent indicator of cellular damage across species (Jin et al., 2011; Zhang et al., 2019; Zahra et al., 2022).

The threat posed by ROS accumulation triggers diverse biochemical defense mechanisms to counteract oxidative stress (Gill and Tuteja, 2010; Fortunato et al., 2023; Li et al., 2023; Nurbekova et al., 2024; Jardim-Messeder et al., 2025; Kračun et al., 2025). While some species enhance their antioxidant capacity, others exhibit reduced efficacy, underscoring the need for detailed analysis of enzymatic and non-enzymatic systems in maintaining cellular homeostasis under heat stress (Foyer and Noctor, 2013; Noctor et al., 2018; Fortunato et al., 2023).

4.2 Antioxidant defense systems

To mitigate ROS-induced damage, plants activate both enzymatic and non-enzymatic antioxidant systems (Foyer and Noctor, 2013; Noctor et al., 2018; Bhuyan et al., 2019; Jardim-Messeder et al., 2025; Jiang et al., 2025; Yang et al., 2025). Key enzymatic antioxidants include superoxide dismutase (SOD), catalase (CAT), peroxidases (POD), and ascorbate peroxidase (APX), which collectively scavenge excess ROS and protect cellular structures (Ru et al., 2023; Jiang et al., 2025; Rao and Zheng, 2025; Zhou et al., 2025). Non-enzymatic antioxidants such as ascorbate and glutathione (GSH) further contribute to redox homeostasis and stress tolerance (Noctor et al., 2018; Jardim-Messeder et al., 2025; Msarie et al., 2025; Rao and Zheng, 2025). The activation of these defense mechanisms varies significantly among plant species (Table 8).

While ROS accumulation under heat stress is well-documented, the efficiency of antioxidant responses shows remarkable interspecific variation (Gill and Tuteja, 2010; Foyer and Noctor, 2013; Noctor et al., 2018; Fortunato et al., 2023; Li et al., 2023; Nurbekova et al., 2024). Some plants exhibit strong activation of both enzymatic and non-enzymatic antioxidants, effectively mitigating oxidative damage (Figure 3), while others show reduced antioxidant capacity, increasing their vulnerability to stress-induced cellular injury (Foyer and Noctor, 2013; Noctor et al., 2018; Bhuyan et al., 2019). These findings reveal a spectrum of defense strategies among species, with some having evolved more effective mechanisms to maintain redox homeostasis. The equilibrium between ROS generation and antioxidant defense is therefore pivotal in determining a plant’s ability to withstand thermal stress and maintain physiological function (Foyer and Noctor, 2013; Noctor et al., 2018). A comprehensive understanding of these biochemical adaptations is essential for developing targeted strategies to enhance crop resilience through breeding and biotechnological approaches.

Figure 3

5 Mitigation strategies against heat stress

Heat stress significantly impairs agricultural crops by disrupting physiological processes and hindering growth and development. Germination rates can decline by up to 95%, while shoot and root growth reductions range from 3.2% to 70% (Tables 1, 2). These morphological changes are accompanied by adaptive physiological and biochemical responses, which may themselves have detrimental effects. To address these challenges, biotechnological and agronomic strategies for enhancing thermotolerance are critical. Promising approaches include the pre-sowing seed priming technique, post-germination supplementary treatments, microbial biostimulation, and microRNA manipulation.

5.1 Pre-sowing seed priming for enhanced plant thermotolerance

Pre-sowing seed priming has emerged as a promising agricultural technique to enhance plant resilience to temperature fluctuations (Filippou et al., 2013; Pagano et al., 2023). This method involves controlled seed treatment to initiate crucial metabolic processes prior to radicle emergence, effectively ‘preparing seeds for faster, more synchronized germination under favorable conditions. Primed plants exhibit superior survival rates under extreme conditions compared to non-primed counterparts (Hönig et al., 2023; Pagano et al., 2023). This method enhances both seed germination and crop productivity through various approaches, including osmopriming, hydropriming, halopriming, and nutrient priming. Moderate temperature exposure has also been shown to boost heat tolerance, primarily by stabilizing the photosynthetic apparatus (Wahid et al., 2008; Ru et al., 2023).

Drought priming, in particular, improves thermotolerance by enhancing morphological parameters and photosynthetic efficiency (Table 9). These benefits are mediated through molecular adaptations, as demonstrated in maize cultivars where thermopriming significantly increases activity of ROS-scavenging enzymes (SOD, APX, POD, CAT), thereby reducing oxidative damage as indicated by lower MDA levels (Ru et al., 2023). Similarly, cold stress priming (0 °C) improves germination rates in barley cultivars (Mei and Song, 2010). The efficacy of priming is closely tied to stress memory mechanisms, where initial mild stress induces epigenetic modifications that enable more robust responses to subsequent stress events (Li and Gong, 2011; Walter et al., 2013; Antoniou et al., 2016). This priming-induced memory represents a powerful tool for developing climate-resilient crops in the face of increasing environmental variability.

H₂O₂ is a cost-effective and widely used priming agent that promotes root/shoot growth and seed germination under heat stress. Its ability to enhance these morphological parameters and germination relies on its capacity to modulate key cellular processes. In maize, H₂O₂ pretreatment enhances photosynthetic efficiency while reducing ROS and MDA concentrations (Wahid et al., 2008). Additionally, it also activates heat stress proteins (27 and 63 kDa) that regulate membrane integrity and structure during root and shoot development, providing molecular support crucial for overall growth (Schoffl et al., 1999; Wahid et al., 2008).

Combined salicylic acid (SA) and chitosan pre-sowing seed treatment enhances rice hybrids’ growth parameters under heat stress, enhancing morphology, photosynthetic performance, and antioxidant defense while reducing ROS and MDA levels (Ahmed et al., 2024). SA alone boosts photosynthetic rate, stomatal conductance, and transpiration rate in rice, though the combined treatment offers superior protection (Ahmed et al., 2024). Notably, unprimed plants are more sensitive to heat during the vegetative stage than during the flowering stages, as seen in gerbera (transpiration rates) and hot pepper (photosynthetic efficiency related to membrane thermostability) (Rajametov et al., 2021; Yang et al., 2023).

SA priming also prevents heat-induced inhibition of RuBisCo, a critical enzyme for carbon fixation, thereby maintaining photosynthetic efficiency comparable to heat-tolerant cultivars (Wang et al., 2010a; Ahmed et al., 2024). By maintaining RuBisCo activity, primed plants exhibit photosynthetic efficiency comparable to heat-tolerant cultivars (Qin et al., 2025; Zheng et al., 2025). This molecular mechanism directly underpins the improved growth and productivity observed in chemically primed plants under heat stress. Likewise, spermidine-treated rice cultivars demonstrate reduced ROS and MDA levels, leading to improved shoot growth and overall plant development under high-temperature conditions (Fu et al., 2019).

Priming activates ROS-scavenging enzymes, heat shock proteins (HSPs), and molecular chaperones, enhancing abiotic stress tolerance in crops like pepper, maize, soybean, spinach, and wheat (Iqbal and Ashraf, 2007; Farooq et al., 2008; Korkmaz and Korkmaz, 2009; Zhuo et al., 2009; Chen et al., 2010). Chemical priming agents, including H₂O₂, abscisic acid (ABA), and SA, improve resilience by modulating photosynthesis, ROS detoxification pathways, and MG metabolism (de Azevedo Neto et al., 2005; Chao et al., 2009; Liu et al., 2010; Wang et al., 2010a, b; Hasanuzzaman et al., 2011; Ishibashi et al., 2011; Gondim et al., 2012; Mostofa and Fujita, 2013; Mostofa et al., 2014; Sathiyaraj et al., 2014; Teng et al., 2014; Ahmed et al., 2024). Hormonal regulation plays a key role, with heat stress increasing ABA while decreasing gibberellic acid (GA) and indole-3-acetic acid (IAA) levels, leading to reproductive impairment (Shimizu and Kuno, 1967; Nakajima et al., 1991; Tang et al., 1996; El-Maarouf-Bouteau et al., 2013; Ishibashi et al., 2015, 2017; Kadyrbaev et al., 2021; Hirayama and Mochida, 2022). These intricate molecular and hormonal interactively collectively contribute to the observable improvements in plant morphology and overall thermotolerance. The synergistic effects of these mechanisms enhance plants’ capacity to maintain physiological function under elevated temperatures. Understanding these hormonal interactions may further refine priming strategies to further enhance plant thermotolerance and improve crop productivity under extreme environmental conditions.

5.2 Post-Germination chemical and nutritional treatments for heat stress mitigation

Beyond pre-sowing seed priming, post-germination treatments provide a valuable complementary approach to enhance thermotolerance in developing plants. The exogenous applications of plant growth regulators, essential nutrients, and protective compounds has demonstrated significant efficacy in maintaining key morphological parameters – including shoot/root growth, floral development, and fruit weight (Zhao et al., 2018; Murad Lima et al., 2025; Ssemugenze et al., 2025; Ali et al., 2021; Khan et al., 2024). While elevated temperatures typically impair photosynthetic efficiency (Tables 5, 6), these supplemental treatments effectively mitigate such physiological limitation during vegetative growth (Table 10). Notably, foliar application of exogenous SA increases chlorophyll content and relative water content (RWC) by approximately 50% in barley while similarly improving growth rates in maize (Khanna et al., 2016; Zahra et al., 2022). The protective mechanisms involve both direct physiological effects and the upregulation of antioxidant defense systems, as evidenced by enhanced activity of ROS-scavenging enzymes (Khanna et al., 2016; Hussain et al., 2019b; Ali et al., 2021; Raghunath et al., 2021; Jahan et al., 2022; Zahra et al., 2022). These coordinated responses collectively strengthen plant resilience to thermal stress while maintaining productivity.

Brassinosteroids post-germination treatments also contribute to heat stress mitigation by promoting reproductive organ growth and enhancing photosynthesis-related markers. In contrast, GA has a stronger impact on morphological traits, such as increasing plant height in rice exposed to 40 °C (Raghunath et al., 2021). Putrescine, a spermidine/spermine precursor, protects plants by regulating potassium channels in guard cells, improving membrane stability. In tomatoes, exogenous putrescine application in the vegetative growth stage activates ROS-scavenging enzymes, increases photosynthetic pigments, suppresses oxidative stress markers, and upregulates HSP 70 and HSP 90 (Jahan et al., 2022).

Melatonin, structurally related to putrescine, enhances shoot/leaf growth while improving tomato thermotolerance when applied as a foliar spray (Khan et al., 2024). Recent studies have highlighted the role of silicon and silicon-containing compounds in stress mitigation as post-germination treatment agents. These molecules stabilize lipid membranes, enhance thermal stability, and promote polysaccharide accumulation, boosting heat resilience (Swain and Rout, 2017; Kumar et al., 2023). Sodium silicate treatment increases barley growth rates by 40% compared to untreated plants (Hussain et al., 2019b).

In addition to these compounds, essential minerals like nitrogen, sodium, and sulfur play a vital role in stress mitigation. Sulfur, crucial for protein biosynthesis and antioxidant defenses, can increase tomato growth rate by 77% under heat stress as it is a central component of thiol groups involved in plant stress responses (Bashir et al., 2015; Ihsan et al., 2019; Ali et al., 2021). While abiotic stress priming and chemical treatments are widely used, biotic stress priming, such as beneficial microbial and viral colonies, also positively influences plant thermotolerance (Khan et al., 2022).

5.3 Microbial biostimulation for heat stress mitigation

The plant microbiome provides additional defense against biotic stressors by enhancing tolerance through a mutualistic relationship (Chauhan et al., 2023; Teiba et al., 2024). Though direct evidence of microbial biostimulation in major crops remains limited, certain bacterial strains support plant growth under heat stress (Table 11). For instance, root-associated bacteria protect tomatoes, while endophytic fungi promote resistance and normal growth (Cameron et al., 2013; Pieterse et al., 2014; Narendra Babu et al., 2015).

One notable example is Paecilomyces formosus, a fungal species that enhances rice growth under normal and high temperatures, likely through secondary metabolite production (Waqas et al., 2015). It also modulates ABA and jasmonic acid (JA) levels, similar to its effects in Capsicum annuum L. and Dichanthelium lanuginosum (Redman et al., 2011; Khan et al., 2015; Waqas et al., 2015). Fungi like Rhizophagus irregularis and Funneliformis mosseae improve maize photosynthetic efficiency (chlorophyll content, fluorescence, rate) while reducing MDA levels through improved N/Mg uptake (Mathur et al., 2021). In tomatoes, Septoglomus constrictum and Septoglomus deserticola enhance morphology and reduce oxidative stress at 42 °C, though without photosynthetic improvements (Duc et al., 2018). Serendipita indica similarly benefits Arabidopsis morphology under heat stress (Chen et al., 2022).

The plant microbiome provides additional defense against biotic stressors by enhancing tolerance through a mutualistic relationship (Chauhan et al., 2023; Teiba et al., 2024). Though direct evidence of microbial biostimulation in major crops remains limited, certain bacterial strains support plant growth under heat stress (Table 11). For instance, root-associated bacteria protect tomatoes, while endophytic fungi promote resistance and normal growth (Cameron et al., 2013; Pieterse et al., 2014; Narendra Babu et al., 2015).

5.4 Mitigation against heat by regulating miRNA

miRNAs (20–24 nt) are emerging tools for improving stress resilience in crops by post-transcriptionally regulating target genes (Chen, 2009; Voinnet, 2009; Asefpour Vakilian, 2020; Ding et al., 2020; Chaudhary et al., 2021; Raza et al., 2021). Furthermore, miRNAs have potential applications in genetic engineering, where their manipulation can improve crop quality and create hybrids with enhanced heat tolerance (Zheng et al., 2025; Zolkiewicz and Gruszka, 2025). Our analysis has identified several miRNAs associated with morphological and physiological changes under heat stress. Moreover, targeted miRNA modifications can contribute to the development of thermotolerant plant hybrids (Table 12).

One key miRNA involved in heat stress response is miR-398, which is upregulated under high-temperature conditions. Under heat stress, miR-398 upregulation suppresses the expression of the copper superoxide dismutase (CSD1/CSD2) while increasing heat shock protein (HSP) expression, improving Arabidopsis thermotolerance (Sunkar and Zhu, 2004; Sunkar et al., 2006; Chen, 2009; Kaczkowski et al., 2009; Guan et al., 2013; Zhao et al., 2016). miR-160 overexpression similarly enhances heat tolerance in Arabidopsis by regulating HSPs and plant development (Lin et al., 2018).

miRNAs also regulate flowering time under heat stress (Li et al., 2022). The transition from the vegetative phase to flowering is controlled by SQUAMOSA PROMOTER BINDING PROTEIN-LIKE (SPL) and APETALA2 (AP2) genes (Stief et al., 2014; Gahlaut et al., 2018; Ma et al., 2020). miR-156 and miR-172 regulate SPL expression, with elevated miR-172 levels (mediated by miR-156) under high-temperature stress leading to reduced SPL expression. This downregulation of SPL subsequently activates FLOWERING LOCUS T (FT), establishing a link between temperature sensing and flowering time (Wu et al., 2009; Kim et al., 2012; Cui et al., 2014; Stief et al., 2014). Additionally, miR-166 overexpression inhibits PHAVOLUTA and REVOLUTA, two genes responsible for leaf development, as well as HOX9, a member of the HD-Zip family (Barik et al., 2014). Similarly, miR-160a downregulates ARF17 and ARF13, affecting the auxin signaling pathway, delaying development, cell proliferation, and flowering (Barik et al., 2014; Kruszka et al., 2014). This delay, together with miR-166’s disruption of leaf development, enhances Arabidopsis thaliana’s tolerance to high temperatures (Reinhart et al., 2002; Mallory et al., 2005; Barik et al., 2014). By precisely regulating stress-response pathways, miRNAs offer powerful biotechnological tools for developing heat-tolerant crops through targeted gene manipulation (Batool et al., 2025; Gao et al., 2025; Spychała et al., 2025; Zheng et al., 2025).

6 Conclusion

Climate change and global population growth pose a severe threat to agricultural sustainability, with rising temperatures particularly diminishing crop yields. High-temperature stress has a negative impact on plant growth and development. While extensive research exists, much of it focuses on individual species or isolated mechanisms, lacking broader comparative insights. This review systematically compared heat stress responses and mitigation strategies across four economically important crops - barley, maize, rice, and tomato - alongside Arabidopsis thaliana. Morphological analysis revealed that root parameters are more sensitive indicators of thermal susceptibility than shoot parameters, though germination rate changes primarily occur under extreme heat. Physiologically, all studied species exhibited consistent reductions in photosynthetic rates and PSII efficiency (Fv/Fm), while stomatal conductance and transpiration rates varied by species and stress duration. Biochemical analyses of ROS accumulation and antioxidant activity presented complex, mixed results - upregulation and downregulation, underscoring the need for deeper investigation into oxidative stress pathways. Among mitigation strategies, priming emerged as an effective one due to its low cost and practical applicability. Other promising approaches include chemical/nutritional treatments, microbial biostimulants, and miRNA-based regulation. Notably, miRNA manipulation offers significant potential for developing thermotolerant crop varieties, promising advancement in both scientific research and agricultural applications.

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