Decrypting molecular mechanism of heat stress tolerance in rice to tackle climate change challenges through recent approaches

Rice (Oryza sativa) is one of the world’s most important cereal crops, contributing to food and financial security, particularly in developing countries. High temperature due to climate change seriously threatens sustainable rice production. Rice crops are adversely affected by heat stress at the morphological, physiological, and molecular levels, resulting in reduced yield and poor grain quality. Rice is highly sensitive to heat during the reproductive phase, causing pollen sterility, impaired pollen dehiscence, pollen germination, and tube growth, ultimately drastically reducing spikelet sterility and yield. High temperature also promotes the accumulation of reactive oxygen species in plant cells, resulting in multiple adverse effects, including damage to chloroplasts and cell membranes, inactivation of photosystems, reduced Rubisco activity, and impaired production of photoassimilates. In this review, we have synthesized the current knowledge on the effects of heat stress on rice and summarized QTLs, genes, and regulatory pathways underlying thermotolerance. We further evaluate conventional breeding, transgenics, and diverse omics-based strategies to breed high-yielding, heat-tolerant rice varieties. The precise molecular insights gained through various omics approaches are expected to advance our understanding of the intricate nature of heat stress tolerance in rice. Additionally, we highlight the emerging roles of microbiome, high-throughput phenotyping technologies, and artificial intelligence as promising tools for accelerating the development of heat-resilient rice.

1 Introduction

Rice (Oryza sativa L.), the most crucial staple food crop supporting over half of the global population, is cultivated across approximately 165 million hectares in 118 countries, with production exceeding 776 million tons in 2022 (FAOSTAT, 2023). Rice production is threatened by earth’s quickly changing ecosystems due to climate change. One of the significant abiotic stresses affecting rice production is high temperature (HT). Since 1850, global temperatures have increased by about 0.06°C per decade (Figure 1A), but warming rate has accelerated sharply to 0.20°C per decade since 1982, over three times faster (NOAA Climate, 2025). The roughly 1°C increase in global average surface temperature since the pre-industrial era (1850-1900), primarily due to greenhouse gas (GHG) emissions into the atmosphere by human activities, might seem small. Still, it means a significant increase in accumulated heat. Historically, the 10 warmest years have all occurred in the past decade (2014-2023). Remarkably, 2023 has been the warmest year by a wide margin. It was 1.18°C above the 20th-century average of 13.9°C. The extra heat leads to regional and seasonal temperature extremes (NOAA, 2024). In the near future (2021-2040), global warming will rise primarily because of the growing cumulative CO₂ emissions in nearly all examined scenarios and modeled pathways. Global warming is more likely than not to reach 1.5°C in the near future even with very low GHG emissions, and is likely to exceed 1.5°C under moderate or high emission scenarios (Lee et al., 2023).

Figure 1

Figure 1. Global warming trends and impact of HS on some rice physiological traits (A) Annual global anomaly (°C) from 1850 to 2023 (Berkeley Earth, 2024), greenhouse gas (GHG) emissions (million metric tons of CO₂ equivalent) (Jones et al., 2024 – with major processing by Our World in Data), and atmospheric CO₂ concentration (ppm) from 1960 to 2023 (NOAA Global Monitoring Laboratory, 2024), illustrating global warming trends. GHGs, particularly CO₂ emissions, are the primary drivers of increased mean air temperature; (B) Effects of HS treatments (control: 25°C, Moderate heat: 35°C, and severe heat: 42°C) on root length, total chlorophyll (Ch) content, and electrolyte leakage (EL) in rice, highlighting physiological responses to elevated temperatures. Increased EL at elevated temperatures indicates membrane damage (Taratima et al., 2022).

1.1 Rising global temperature and its effect on rice production

Prolonged exposure to temperatures surpassing a critical threshold (Table 1) conducive to optimal physiological functions inflict irreversible damage on plant growth (Khan et al., 2019). It is particularly significant in Southeast Asia, where rice plays a vital role, providing 3/4^th of the region’s calorific intake (Fitzgerald et al., 2009). However, this region is predicted to experience the most rapid temperature increase (IPCC, 2014). By 2030, it is extrapolated (Gourdji et al., 2013) that approximately 16% of the rice growing areas will encounter at least five reproductive days with temperatures surpassing Tcrit (physiological critical temperatures during the reproductive stage). This proportion is anticipated to rise to 27% by 2050. Between 2022 and 2023, a noticeable decline in global rice production, amounting to an estimated reduction of 300,000 tons compared to the preceding year, 2021-2022, was seen (USDA 2023). Notably, while specific rice-producing countries like India, Thailand, and Türkiye managed to sustain their production, several other prominent rice-producing countries have encountered considerable yield losses in recent years, attributed to the impacts of climate change. The trend is exacerbated by the fact that the world’s population is growing at a rate of 0.88% annually, necessitating expedited efforts to enhance rice productivity to keep pace with the demographic demand (UN DESA, 2022).

Table 1

Table 1. Critical temperatures for the development of the rice plant at different growth stages.

According to (Peng et al., 2004), research from the International Rice Research Institute (IRRI) during 1992–2003 indicates yield drop in rice by 10% for every 1°C rise in minimum temperature during the growing season. Similarly, a daytime temperature increase of 28°C to 34°C decreases the yield by up to 7-8% (Korres et al., 2017). A heatwave in Japan led to a 25% spikelet sterility rate when temperatures peaked around 38°C in 2007 (Hasegawa et al., 2011).

1.2 Projected declines in rice production due to elevated temperatures

Population growth has created a critical demand to ramp up crop production for food security. Estimates suggest that a 70% boost in food production will be vital to cater the demands of an anticipated 9 billion population by 2050 (Bita and Gerats, 2013). Short-term projections indicate that rice production in South Asia could decline by about 10% by 2030 (Lobell et al., 2008). Medium and long-term estimates predicts a 10-25% reduction in rice yields across developing countries by 2080, with India potentially facing losses of 30-40% due to extreme heat events (Cline, 2007). Overall, high temperature stress (HTS) may lower rice grain yield by up to 41% by the end of the 21^st century, as the temperatures are expected to rise by up to 2°C by 2050 relative to 1950 (Ceccarelli et al., 2010). Without the benefits of CO₂ fertilization, effective adaptation measures, and genetic improvement, each 1°C rise in global average temperature is projected to decrease worldwide wheat yields by 6.0%, rice yields by 3.2%, maize yields by 7.4%, and soybean yields by 3.1% (Zhao et al., 2017). Spatial modeling predicts a 20% decrease in boro rice yield in Bangladesh by 2050, escalating to 50% by 2070, with average rice yields declining by up to 33% by 2081–2100 (Basak et al., 2009; Karim et al., 2012).

2 Heat stress combined with drought: amplifying stress responses in rice

Although rice is susceptible to heat and drought (Kumar et al., 2014; Venuprasad et al., 2007), the combination of drought and heat stress (HS) is the most common abiotic stress in field conditions, significantly impacting crop productivity. The simultaneous occurrence of drought and HS in various rice-growing regions is almost inevitable, leading to increased plant-tissue temperature as drought severity progresses. Mechanistic studies have shown that the combined exposure to drought and HS elicits a unique response rather than a simple additive effect of both stresses (Rizhsky et al., 2002; 2004). Despite recognizing the practical importance of combined drought and HS on plants, there is limited field-based knowledge in this area (Lawas et al., 2018). While the effects of combined heat and drought stress have been studied in model plants, relatively little information is available on rice’s response to these stresses, particularly during the critical flowering stage (Rang et al., 2011). Understanding the molecular mechanisms of tolerance to this stress combination during sensitive flowering and grain-filling stages in cereals, especially rice, remains limited (Lawas et al., 2019). Empirical screening for thermotolerance at different stages, and evaluating heat tolerance under combined stress conditions could accelerate the development of rice varieties with improved tolerance to multiple stresses (Costa et al., 2021).

2.1 Types of stress responses to heat in rice plants

Understanding the mechanisms by which rice plants respond to elevated temperatures is crucial for answering the key question: how do rice plants sense HT and then transduce signals into intracellular responses? This knowledge is equally critical to breeding rice cultivars with improved HS tolerance. Three types of different plant responses have been observed under HTS, namely basal thermotolerance, acquired thermotolerance (AT), and programmed cell death (PCD) (Guihur et al., 2021; Haider et al., 2021; Locato and Gara, 2018; Mittler et al., 2012). Basal thermotolerance is an inherent ability to survive HT above those conducive for growth and to acquire tolerance to lethal temperatures. In contrast, acquired thermotolerance, which is also known as adaptive thermotolerance, is the ability to withstand an otherwise lethal HT after being pre-exposed to a sublethal increased temperature, mimicking an ‘immunization’ against HT (Larkindale et al., 2005; Lim et al., 2013; Shanmugavadivel et al., 2019). Plants may remove some specific cells in response to HT or other environmental stimuli in a process called PCD (Locato and Gara, 2018).

3 Necessity for thermotolerance rice breeding

The need to breed heat-tolerant rice is crucial, considering its critical role in global food security and the detrimental effects of climate change on yield. To breed rice with heat tolerance, it is essential to elucidate the molecular basis of HS response in rice, the genes, proteins, and physiological and biochemical traits that confer heat tolerance (Janni et al., 2020; Raza et al., 2020; Sailaja et al., 2015). This review provides an overview of HS-induced morphological and physiological changes, elucidating molecular mechanisms underlying the HS response regulatory network in rice and strategies to breed for enhanced rice adaptation to global warming through various approaches.

4 Stage-specific effects of heat stress on rice

4.1 Effect of HTS on germination and vegetative growth

Each stage of rice plant development responds differently to HT (Zhang et al., 2018). HS has an impact on grain quality, dormancy, germination, and emergence in addition to seedling vigor and establishment across the entire seed development process (Brunel-Muguet et al., 2015; Finkelstein et al., 2008; Liu et al., 2019). Exposure to HT during seed germination leads to lower germination rates and decreased vigor in germinated seedlings (Fahad et al., 2017; Liu et al., 2019) (Figure 2). At the seedling stage, rice grows best at a temperature between 25 and 28°C. In seedlings, elevated temperatures (42–45°C) (Table 1) can damage cell membranes, hinder photosynthesis, and escalate oxidative damage, which results in increased water loss, wilting, impaired root growth (Figure 1B), and potentially plant death (Bahuguna et al., 2015; Liu et al., 2018). A decrease in germination and seed vigor due to HS has been associated with reduced plasma membrane (PM) thermostability and membrane fluidity (Fahad et al., 2017; Saidi et al., 2010). Lipid profiles of PM acclimatized to moderate HS revealed a marked reduction in fatty acid unsaturation, leading to increased membrane rigidity. This structural change accounts for the attenuated Ca²⁺ influx observed during HS (Saidi et al., 2010; Sangwan et al., 2002). Begcy et al. (2018) reported that HS (35°C) dramatically decreases the size of grain at maturity because of lower length, breadth, and mature grain weight during early grain development; and when the temperature reaches 39°C, the endosperm collapses, and seed viability is significantly reduced. Tillering, a crucial agronomic trait in rice, is severely affected by HT and thus reduces the number of panicles per plant. Soda observed a 35% reduction in panicle number and a 28% decrease in yield per plant in rice plants exposed to elevated temperatures (Soda et al., 2018). Other morphological traits to assess under HS include leaf drooping and rolling, reductions in plant biomass, and decreased chlorophyll concentration (Ali et al., 2022; Ren et al., 2023; Sita et al., 2017).

Figure 2

Figure 2. Effect of HS on rice at different stages. (A) Vegetative stage: HS during seed germination leads to low germination rates and week seedling vigor. Tiller numbers are reduced due to disrupted cell membranes, impaired photosynthesis, and accumulation of ROS. Phytohormone imbalances, increased water loss due to increased stomatal density, wilting, and impaired root growth further negatively impact the development of rice plants. (B) Reproductive stage: HS leads to altered flowering time, deformed floral organs, reduced pollen-viability, -germination and -tubegrowth, and spikelet sterility. Inhibited pollen dehiscence affects the amount of pollen landing on a stigma and negatively impacts fertilization rates. (C) Ripening stage: Decreased photosynthetic rate causes inhibited assimilate production and accelerates the senescence of functional leaves, resulting in reduced partitioning of carbohydrates into the grains. During the milky stage, HS hinders the synthesis and movement of carbohydrates, proteins, and lipids in developing grains. The initiation of endosperm cellularization is a critical developmental transition required for normal seed development. Impaired cellularization hinders endosperm development. Schematic drawing shows early stages of endosperm development (ED) under control and severe HS conditions. Under normal conditions, the rice seed development follows syncytial ED, early cellularization, and mid-cellularization stages. Under HS conditions, initiation of cellularization is severely affected. The central vacuole (CV) remains present when seeds are exposed to severe HS.

Rice seedlings’ ability to withstand HT varies depending on their genetic composition. The domestication origins of the two subspecies, japonica and indica, differ. The japonica emerged in the temperate regions, while the indica originated in tropical areas. The indica exhibits greater thermotolerance than japonica and possesses distinct morphological and physiological traits (Lee, 2002; Lee et al., 2017). HS affects tiller and panicle numbers more significantly in Japonica rice relative to Indica rice (Wang et al., 2016). Regarding heat resistance, hybrid rice varieties combining indica and japonica demonstrate the highest level, followed by indica and then japonica varieties individually (Prasanth et al., 2017).

4.2 Impact of HTS on the reproductive stage

The reproductive phase (panicle initiation to physiological grain maturity) is the most vulnerable stage to abiotic stresses (Guan-fu et al., 2008). The stages of panicle initiation, formation of male and female gametophytes, anthesis, pollination, and fertilization are most vulnerable to HS (Arshad et al., 2017; Jagadish et al., 2015). According to Xu et al. (2020), both daytime and nighttime HS causes deformation of floral organs reducing their size and number. During anthesis, HS impairs pollination, significantly increasing spikelet sterility (Sarangthem et al., 2021). Reactive oxygen species (ROS) is accumulated in plant cells during HT, resulting in multiple adverse effects such as damage to the chloroplast and cell membranes, loss of activity of photosystems, suppressed RuBisCo activity, and decreased production of photoassimilates. These issues culminate in poor flowering and decreased grain yield (Lal et al., 2022; Radha et al., 2022; Zaidi et al., 2019). HT has multiple adverse effects on rice stamens as (I) elevated temperatures disrupt meiosis during the pollen development, disintegration of tapetum and/or reduced activity of invertase enzyme, leading to the production of sterile pollen (Endo et al., 2009) (Figure 3), (II) HT inhibit pollen dehiscence and reduce the swelling capacity of pollen grains, which diminishes pollen amount landing on a stigma and negatively impacts fertilization rates (Arshad et al., 2017; Hu et al., 2021), (III) the moisture content adjusts of the pollen grains is essential for their formation and dispersion. The pollens landing on stigma adjust their moisture levels to environmental conditions, but HT can disrupt this process (Das et al., 2014; Shrestha et al., 2022), (IV) HT significantly decreases the protein content in the pollen, decreasing its germination ability and pollen-tube elongation rates, which ultimately leads to spikelet sterility (Arshad et al., 2017; Jagadish, 2020; Shrestha et al., 2022). HTS during anther formation, particularly during pollen mother cell meiosis, can lead to early deterioration and breakup of tapetal cells. This affects the nutrition of microspores and the generation of pollen walls, culminating in abortion of pollen grains and reduced stigma length (Liu et al., 2020; Xu et al., 2020; Zhang et al., 2018) (Figure 3). An indica variety, IR64, showed 66% reduction in the number of spikelets when it was exposed to HS (40°C day/35°C night) at pre-flowering stage to HS conditions for 15 days (Soda et al., 2018). Hu et al. (2021) observed a decline in pollen viability, spikelet fertility, and grain yield by 46%, 69%, and 84%, respectively under HS in a heat susceptible variety Liangyoupeijiu (LYPJ) in comparison to 18% yield reduction in Shanyou63 (SY63), a heat tolerant variety. Lin et al. (2023) found that heat-sensitive rice mutant, heat shock protein60-3b (oshsp60-3b), showed decreasing fertility as temperature increases. Overexpression of OsHSP60-3B enhanced thermotolerance of pollen in transgenic plants. Multiple genes regulating heading in rice such as Hd1 (heading date 1), Ehd1 (early heading date 1), Ghd7 (grain number, plant height, and heading date 7), and Hd3a/RFT1 (heading date 3a/rice flowering locus T1) form the core Ghd7-Ehd1-Hd3a/RFT1 flowering pathway. HT reduces GhD7 transcript levels, reducing its inhibitory effect on RFT1 and enabling timely floral induction. A natural allele of qHd1 (encoding OsMADS51) further enhances heat tolerance at heading and grain filling through OsMADS51-Ehd1-Hd31/RFT1 pathway (Kan et al., 2023; Zhou et al., 2021; Xue et al., 2008).

Figure 3

Figure 3. Impact of HS on the reproductive phase. (A) Impaired male and female gametophyte development: Floral organs are deformed under HS and have reduced numbers and sizes. HS leads to significant changes in gene expression, resulting in tapetum degeneration and pollen sterility in developing anthers during the early microspore stage. Swelling of pollen grains is restricted at the time of heading under HS conditions. Pollen fertility also decreases due to delays in the opening of the locule. HS severely reduces anther dehiscence during anthesis. Pollen viability and germination are reduced due to decreased protein and iron content in pollen grains. Pollen tube growth is significantly affected by HS. (B) Male gametophyte development under normal conditions. (C) Male gametophyte development under HS conditions: Developing pollen and the surrounding tapetal cells are highly sensitive to HS, resulting in premature degeneration of tapetal cells, causing disruptions in the supply of nutrients to developing pollens. HS affects the meiotic cell division by influencing the orientation of the spindle apparatus, resulting in aberrant chromosome behavior and failure of pollen development. ROS accumulation is increased, and soluble carbohydrate and starch reserves are decreased in developing anthers under HS, leading to starved microspores and increased pollen sterility.

In addition, HT severely impact the further fertilization processes. Restricted pollen tube growth hinders the pollen movement towards egg cells because of disruption in the ion balance, carbohydrate metabolism, and phytohormone concentration of pollens (Coast et al., 2016; Firon et al., 2006; Jagadish, 2020; Yan et al., 2002). Following double fertilization, a short term exposure to HTS (39°C for 48 hours) leads to impaired cellularization during initial endosperm development, hindering the subsequent establishment of the endosperm (Folsom et al., 2014). Many studies in recent years have been carried out to investigate the effect of HS on the morphology of the reproductive parts. Still, fewer reports are available on the impact of HS on stigma. Jagadish et al. (2010) observed reduced stigma length when they exposed the rice plants to HT for 6 hours during anthesis. Increased stigma length may enhance tolerance to HS during the flowering period. Callose could be used as an indicator of sterile ovules, with its deposition at the ovule chalaza commonly used to assess early ovule degeneration (Endo et al., 2009). Evaluating and selecting rice varieties on the basis of characteristics such as rapid pollen dehiscence (Kobayashi et al., 2011), proper septum breakage during pollen expansion (Matsui and Omasa, 2002), and increased pollen protein content (Arshad et al., 2017) can provide more comprehensive insights into pollen quality and offer superior indicators than pollen viability alone when screening for HS tolerance. Additionally varieties with enhanced internal anthocyanin concentration have been shown to protect the photosynthetic apparatus via ROS scavenging mechanism, thereby improving thermotolerance (Zaidi et al., 2019). Early-flowering rice varieties, which can escape HTS, are also considered cost-effective and widely adopted in plant breeding. The QTL qEMF3, detected in Oryza officinalis, shifts the flower opening time of cultivars to earlier in the morning (Hirabayashi et al., 2015; Jagadish, 2020; Jagadish et al., 2007). Substantial changes in the metabolic profiles of different tissues in rice are observed under HS conditions. Specifically, heat-tolerant rice varieties exhibit a unique build-up of crucial metabolites, setting them apart from heat-sensitive types (Singh et al., 2024) (Figure 4). Consequently, analyzing the morphology and physiology of flowers in various heat-tolerant and -sensitive varieties could assist plant breeders in identifying and selecting for thermotolerance.

Figure 4

Figure 4. Metabolic profiling of pistils at the end of HS in N22 and IR64. Metabolite profiling of pistils in varieties N22 (heat tolerant) and IR64 (heat sensitive) after HS temperature treatment reveals the distributions of metabolites under control (30°C) and HS conditions (38°C and 40°C). Metabolites involved in significantly altered metabolic pathways are shown here. Under control conditions, IR64 showed higher concentrations of most of the metabolites compared to N22, except for glycine and asparagine, which were significantly lower in IR64. After HS treatment in N22, all the metabolites from the TCA cycle (pyruvic acid, citric acid, oxoglutaric acid, succinic acid, fumaric acid) increased under 38 °C and 40 °C compared to the control, except for oxoglutaric acid at 38 °C (Shi et al., 2022). "*" represents significant difference between genotype metabolites for different temperature treatments.

4.3 Impact of HTS on rice grain development and quality

Elevated temperatures hinder the transport and biosynthesis of sugars, proteins, and lipids in rice caryopses during the milky stage (Figure 2), consequently affecting grain weight and quality (Liao et al., 2012; Sreenivasulu et al., 2015; Zhang et al., 2016). An enhanced rate of grain filling and a reduced total grain filling duration of 21.3%–37.1% for various genotypes after HS treatment at the grain-filling stage have been reported. After being subjected to HT (38 °C/30 °C day/night) continuously for 20 days at grain filling stage, the seed weight decreased by 24.6% for tolerant Nagina 22 (N22) and 39.1% for sensitive IR64 compared to normal conditions (31 °C/23 °C day/night) (Shi et al., 2017; Zhang et al., 2022). Grain filling is more prone to high night temperature (HNT) stress (Krishnan et al., 2011). HT inhibits assimilate synthesis by decreasing photosynthesis (Zhang et al., 2009) and accelerating the senescence of functional leaves, resulting in reduced assimilate transfer to grains. Additionally, elevated temperatures may hinder early embryo (Cao et al., 2016) and seed formation (Huang et al., 2019). HT can supress gene expression and often disrupt the bioactivity of starch-producing enzymes, thus disturbing starch accumulation and the ratio of amylose to amylopectin in the endosperm of rice (Chen et al., 2017; Impa et al., 2021; Yamakawa and Hakata, 2010; Zhang et al., 2021). HS has also resulted in lower grain weight, decreased grain width, altered kernel size, and reduced yield (Folsom et al., 2014; Lyman et al., 2013; Rangappa et al., 2024; Shi et al., 2017) (Figure 5). HT at grain filling stage has shown to cause a significant decrease in rice yield, with losses of up to 50% (Sreenivasulu et al., 2015). Under HS conditions, a decrease of 16.7% in number of grains per plant led to a significant reduction of 18% in grain yield at HNT (28°C from 6 pm to 6 am) (Sharma et al., 2024).

Figure 5

Figure 5. Impact of HS on different yield parameters of various rice varieties. Different popular rice varieties show a wide range of variation compared with heat-tolerant varieties (N22 and NERICA-L-44) under HS conditions for single plant yield (grams), test weight (grams), number of effective tillers, and spikelet fertility percentage (Surender et al., 2021).

In addition to affecting yield, rice grain quality is also compromised. It has been demonstrated that HT during the grain filling period increases the chalkiness rate and decreases head rice rate, gel consistency, amylose content, and overall taste quality (Dou et al., 2024). Setback viscosity, average particle size, crystallinity, and gelatinization temperature increased when the plants were subjected to 34°C (5°C higher than the normal temperature) (Zang et al., 2022). HTS accelerate non-uniform grain filling due to rapid endosperm cell division, ultimately shortening the filling period. Poor starch accumulation pattern and loose packing of starch granules results in chalkiness, brittleness, fissured, and broken grains. These reduces palatability, appearance, and milling quality of rice (Sreenivasulu et al., 2015; Yao et al., 2020; Shirdelmoghanloo et al., 2022; Zhang et al., 2021; Nevame et al., 2018). When plants were subjected to HS during the grain filling stage at 38/30°C (day/night) temperature, increased chalkiness (averaging 196.6%), reduced length/width ratio, increase in protein content (7.8 – 29%), decrease in total starch content (1.5%), decrease in amylose (6 - 11.5%) and decrease in amylopectin (5.2%) were observed (Zhang et al., 2023).

Starch in rice grains is the primary reserve, and its biosynthesis is very sensitive to heat. Among the various enzymes contributing to starch biosynthesis, ADP-glucose pyrophosphorylase (AGPase) is particularly susceptible to HS in the seed maturation phase. To increase thermotolerance for this enzyme in rice, Hwang et al. (2019), manipulated two dominant subunits of AGPase present in the developing endosperm, the large (L2) and small (S2b) subunits of cytosol-specific AGPase.

5 Impact of HTS on rice physiology

5.1 Impact of HTS on membranes

Plant cells’ primary protective barriers are biomembranes, composed of highly organized lipids and proteins, that are extremely sensitive to heat (Niu and Xiang, 2018; Sita et al., 2017). Impaired plant growth and development are often associated with plant cells’ disrupted physiological and metabolic processes. The increased kinetic energy and protein mobility stimulated by elevated temperatures lead to molecular bonds within membranes (Dhanda and Munjal, 2012). HST can disrupt the composition and function of PM, altering the ratio of saturated to unsaturated fatty acids and inducing protein denaturation. This disruption leads to elevated fluidity and permeability, impaired membrane integrity, and higher leakage of cells’ ions (Xalxo et al., 2020). The primary response to elevated temperatures also occurs in the PM, which activates the channels and receptors essential for HS sensing, cellular response, and transduction of calcium signalling. HTS also hampers the activity of fatty acid desaturases, hence affecting the extent of unsaturation of fatty acid chains, crucial for HS adaptation in plant (Higashi and Saito, 2019; Lamers et al., 2020; Niu and Xiang, 2018; Shen et al., 2015).

To survive extreme temperatures, plant cells must first ensure the stability and fluidity of their lipid membranes by modulating lipid saturation, with fatty acids being the fundamental components. HTS1 is crucial for de novo fatty acid biosynthesis, and its scarcity inhibits fatty acid synthesis and metabolism of fats in hts1 mutants. This reduction in fatty acid level compromises cell membrane’s integrity and stability under HS, which leads to abnormal heat-induced calcium signaling (Chen et al., 2021). OsCNGC14 and OsCNGC16, which are cyclic nucleotide-gated ion channels located in the PM, regulate calcium signals in response to HTS, thereby conferring thermotolerance to rice (Cui et al., 2020).

Membrane thermostability is the most reliable trait for screening heat-tolerant rice genotypes, showing a strong correlation with yield under HT. Mean relative injury and leaf electrolyte leakage are increased by HTS (Figure 1B). Therefore, genotypes with high membrane thermostability and low relative injury are promising candidates for direct selection or hybridization in future breeding programs for rice thermotolerance (Sailaja et al., 2015; Maavimani et al., 2014).

5.2 HTS induced alterations in photosynthesis

HS disrupts the permeability of the thylakoid membrane and even cause the disintegration of thylakoid grana, which leads reduced in chlorophyll (Figure 1B) and altering photochemical reactions, causing a reduction in ratio of variable fluorescence to maximum fluorescence (Fv/Fm) and rate of photosynthesis (Chakraborty and Bhattacharjee, 2015; Hu et al., 2020; Wang et al., 2018). Among these components, photosystem II (PS II) is particularly susceptible to HS, with oxidative stress induced by the HT causing the dissociation of the oxygen-evolving complex (OEC) in PSII, thereby restricting electron transport from OEC towards the acceptor side of PSII, thus causing a substantial decline or complete loss in its activity (Essemine et al., 2017; Sailaja et al., 2015; Szymańska et al., 2017). Moreover, HT inhibits the activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCo), primarily due to the inactivation of RuBisCo activase (Perdomo et al., 2017). Other than this, HT also reduces amounts of photosynthetic pigments and leads to reduced carbon fixation potential (Hasanuzzaman et al., 2013; Song et al., 2014).

Genetically engineered transgenic plants having increased levels of RuBisCo activase showed improved growth in HT conditions and demonstrated higher photosynthetic rates compared to wild-type plants (Wang et al., 2010). Overexpression of a thermotolerant RuBisCo activase from wild rice significantly enhanced growth and grain yield of cultivated rice during HT, indicating that manipulating RuBisCo activase could be an efficient strategy for thermotolerance rice breeding (Scafaro et al., 2018). To prevent damage buildup, PSII plants utilize the de novo synthesis of proteins, including the D1 subunit protein (core subunit of PSII, susceptible to light and HS) encoded by the chloroplast gene psbA, which is crucial for the process. Introducing a heat-responsive promoter to enhance D1 protein expression increased heat tolerance, resulting in significant increases in both aboveground biomass (20.6%–22.9%) and grain yield per plant (8.1%–21.0%) compared to regular rice plants (Chen et al., 2020). The absence of OsNSUN2 (an RNA 5-methylcytosine (m⁵C) methyltransferase) function results in a weakened photosystem characterized by decreased efficiency in photosynthesis and the accumulation of ROS following exposure to heat (Tang et al., 2020). Ahmad et al. (2024) observed that the gene PALE GREEN LEAF 10 (PGL10) is essential for chlorophyll synthesis in rice. Loss of PGL10 function results in pale green leaves and impaired photosynthesis under HT conditions. Thus, thoroughly understanding how photosynthetic metabolism responds to HS is essential for examining plant resilience and recognizing the detrimental effects of HT on agricultural productivity (Bita and Gerats, 2013).

5.3 Impact of HTS on carbohydrate metabolism and partitioning

HTS alters carbohydrate metabolism and the distribution of photo-assimilates in rice plants (Arshad et al., 2017; Bahuguna et al., 2016; Shi et al., 2017). The levels of two crucial enzymes involved in the glycolytic pathway, phosphoglucose isomerase and phosphofructokinase, were significantly reduced, along with a decrease in the abundance of phosphoglycerate mutase, when rice cells were subjected to HT (44°C). This suggests that a cell’s ability to produce energy under HS is hampered (Gammulla et al., 2010). Compared to wild-type plants, a heat-resistant rice mutant, ett1, demonstrated increased survival, less oxidative damage, and higher photosynthetic efficiency under HTS. This mutant’s accumulation of higher energy and carbohydrates suggests enhanced metabolic activity and stress tolerance (Feng et al., 2023). A transcription factor (TF), HYR (higher yield rice) involved in carbon metabolism, enhances photosynthesis under HS (Ambavaram et al., 2014).

HTS disturbs sugar content in anthers, disrupting the regular nutrient supply essential for developing pollens (De Storme and Geelen, 2014; Rezaul et al., 2019). In rice varieties sensitive to heat, the Carbon-Starved Anthers (CSA) gene has increased expression. In contrast, heat-resistant varieties show robust expression of the sugar transporter gene MST8 and the cell wall invertase gene INV4. This suggests that sugar deficiency has a significant function in spikelet sterility (Li et al., 2015). In a similar vein, compared to a sensitive cultivar, the thermotolerant rice cultivar expressed more of the sucrose transporter gene OsSUT1, which increased the amount of photo-assimilates available for filling kernels (Miyazaki et al., 2013).

5.4 Impact of HTS on phytohormones

Plant hormones are vital for regulating growth and development of rice plants under optimal and adverse environments by triggering many signaling cascades to facilitate the adaptive responses of plant (Khan et al., 2023). Exogenous application of various compounds, including antioxidants (ascorbic acid, alpha-tocopherol), amino acids (glycine betaine), and plant hormones (salicylic acid, auxins, brassinosteroids, methyl jasmonates) has highlighted the significant role of phytohormones in mitigating the adverse effects of HTS on rice (Fahad et al., 2016; Mohammed and Tarpley, 2011). During HTS, salicylic acid has been found to reduce the accumulation of ROS in anthers to mitigate pollen abortion by preventing premature degradation resulting from tapetum programmed cell death (Feng et al., 2018; Nadarajah et al., 2021). Melatonin deficient rice mutants coupled with reduced brassinosteroids (BRs) synthesis exhibited enhanced tolerance to HS (Hwang and Back, 2019). Ethylene-mediated signaling pathways aid in minimizing oxidative damage, preserving chlorophyll content, and regulating metabolism of carbohydrates which leads to improved thermotolerance in seedlings (Gautam et al., 2022). HS alters the balance of phytohormones, causing decreased active cytokinin (CTK), gibberellin (GA), and indole-3-acetic acid (IAA) contents in rice spikelets and developing kernels. This disruption hinders cell proliferation and panicle formation, decreasing spikelet number, pollen fertility, and kernel weight. Conversely, HS results in elevated abscisic acid (ABA) content in anthers and seeds, which, as a result, induces pollen abortion and inhibits germination and seedling establishment (Liu et al., 2019; Tang et al., 2008; Wu et al., 2016). Exposure to HS inhibits CTK transportation rate and CTK synthesis enzymes while increasing cytokinin oxidase/dehydrogenase activity, particularly in heat-sensitive cultivars. These changes likely contribute to decreased panicle CTK abundance under HS conditions (Wu et al., 2017).

Zhang et al. (2018) studied how levels of naturally occurring ascorbic acid affect the use of transgenic rice plants. These plants either had increased or decreased activity of L-galactono-1,4-lactono dehydrogenase (GLDH) enzyme, which catalyzes the terminal step in ascorbic acid biosynthesis pathway. When GLDH activity was suppressed, higher levels of ROS were observed in the transgenic rice. However, when endogenous ascorbic acid was present at higher levels, it inhibited the breakdown of RuBisCo and chlorophyll. Consequently, it reduced ROS accumulation, enhancing rice plants’ stability when exposed to HT. OsNCED1 (9 CIS-EPOXYCAROTENOID DIOXYGENASE) can control the endogenous ABA content in rice. This enzyme enhances the antioxidant capacity, activates the expression of gene associated with heat and ABA, and positively regulates rice seedling thermotolerance (Zhang et al., 2022). By upregulating two heat-responsive genes, JASMONATE ZIM DOMAIN (JAZ), and the heat tolerance gene on chromosome 3 (HTG3) controls rice thermotolerance (Wu et al., 2022).

Thus, comprehending how these plant hormones function will significantly aid in identifying the mechanisms of HT tolerance in rice plants. To mitigate the impact of HS on yield losses, strategies could involve manipulating phytohormone signaling pathways to develop varieties with enhanced heat tolerance.

5.5 Impact of HTS on ROS accumulation

Studies have shown that the exposure to HTS induces a rapid ROS burst in plant tissues and disrupts the homeostasis between ROS production and detoxification (Baxter et al., 2014; Huang et al., 2016; Zhao et al., 2023). ROS affects the redox homeostasis and the function of proteins including their transcriptional activities and enzymatic properties during stress situations (Mittler, 2017). HT triggers the production of hydrogen peroxide (H₂O₂) primarily in chloroplasts and mitochondria. This compound not only serves as an early messenger in cellular signaling but also inflicts damage on cells over time, potentially leading to programmed cell death (Qiao et al., 2015), growth retardation, and grain chalkiness (Suriyasak et al., 2017), seedling death (Fang et al., 2015) and spikelet sterility. In rice plants experiencing HTS (with days at 38 °C and nights at 30°C during meiosis), the ROS concentration in anthers exceeds threefold that of normal temperature conditions (with days at 28 °C and nights at 22°C). HTS also induces a surge in ROS levels in rice pistils, likely due to the upregulation of Respiratory Burst Oxidase Homolog (RBOH) genes (Fu et al., 2016; Zhang et al., 2018, 2023). Additionally, HTS hampers the activity of antioxidant enzymes, particularly superoxide dismutase (SOD) and catalase (CAT) (Sailaja et al., 2015; Zhang et al., 2018).

OsANN1, an annexin that binds calcium, enhances thermotolerance through modulating antioxidants accumulation such as CAT and SOD under HT (Qiao et al., 2015). Excessive ROS, particularly, exacerbates membrane lipid peroxidation and protein oxidation, resulting in heightened levels of intracellular malondialdehyde (MDA), which can disrupt the normal functioning of proteins and nucleic acids (Bahuguna et al., 2015; Chakraborty and Bhattacharjee, 2015). Therefore, indicators such as electrolyte leakage, ROS levels, expression levels of antioxidative genes, activities of antioxidant enzymes, and MDA content are regularly employed to assess membrane and oxidative damage and measure plants’ heat tolerance. For instance, heat-tolerant rice varieties like NERICA-L-44 and N22 demonstrate increased membrane stability and reduced ROS and MDA levels due to elevated antioxidant enzyme activities (Bahuguna et al., 2015; Higashi and Saito, 2019; Sailaja et al., 2015). Heat-sensitive mutants accumulated ROS, reduced catalase activity, and upregulated OsSRFP1 (a RING finger E3 ubiquitin ligase) expression under HS (Zafar et al., 2020). Overexpressing OsProDH (encoding protein dehydrogenase) lines had reduced proline content while its knockout mutant lines had enhanced proline content. Increased proline content reduced H₂O₂ accumulation in the seedlings of mutant lines indicating OSProDH regulating HS tolerance negatively (Guo et al., 2020).

Monodehydroascorbate reductase (MDHAR), encoded by OsMDHAR4, is a scavenger of ROS. OsMDHAR4 negatively impacts rice thermotolerance by modulating stomatal responses induced by H₂O₂. Inhibition of OsMDHAR4 promotes stomatal closure, increases H₂O₂ accumulation, reduces water loss, and enhances heat tolerance (Liu et al., 2018). The pyridoxal phosphate homeostasis protein (PLPHP), encoded by HTH5 and mitochondria-localized, may mitigate damage to mitochondrial energy metabolism during HT by controlling ROS dynamics. Overexpressing HTH5 significantly mitigates ROS accumulation triggered by HS (Cao et al., 2022).

6 Impact of HTS on protein homeostasis

Another significant consequence of HS is the perturbation of protein homeostasis or proteostasis within cells, resulting in cell death and toxicity. The term proteostasis includes processes associated with biogenesis, folding, unfolding, trafficking, and turnover of proteins (Mishra and Grover, 2016). HS often causes protein misfolding, unfolding, and protein denaturation or aggregation in the endoplasmic reticulum (ER) and triggers the unfolded protein response (UPR) to restore ER protein homeostasis (Liu and Howell, 2016; Sun et al., 2021; Zhang et al., 2017). A group of specialized proteins known as Heat Shock Proteins (HSPs) function as chaperones and can be crucial in stabilizing, correcting, refolding, restructuring, compartmentalizing, or breaking down misfolded proteins (Sharma et al., 2019). OsHSP101 has been observed to perform a key function in enhancing the long-term acquired thermotolerance in rice by forming a positive feedback loop with HSA32(HEAT STRESS ASSOCIATED 32-KD PROTEIN) (Lin et al., 2014). When toxic proteins accumulate rapidly, proteasome-mediated degradation proves more effective than HSPs in restoring the denatured proteins. Specifically, TT1 (Thermotolerance1), an α2 subunit of the 26S proteasome, efficiently removes cytotoxic denatured proteins associated with ubiquitination, helping to maintain protein homeostasis during HS. Overexpression of OgTT1 markedly increased thermotolerance in rice, Arabidopsis, and Festuca elata (Kan and Lin, 2021; Li et al., 2015).

7 Impact of HTS on source and sink dynamics

During HTS plants experience cellular disruption due to redox imbalance and increased ROS leading to reduced photosynthetic efficiency and impaired activities of enzymes associated with sugar metabolism, ultimately affecting source-sink dynamics (Gautam et al., 2022; Hassan et al., 2020). Grain filling consists of using carbohydrates like sucrose, made in the leaves, i.e., source. These sugars travel long distances through the phloem, which acts as a transport pathway, and are finally deposited into the developing endosperm, where they are stored as starch granules in amyloplasts (Nazir et al., 2023; Ren et al., 2023).

GRAIN FILLING RATE 1 (GFR1) boosts sucrose production in leaves by enhancing RuBisCo activity, consequently regulating the grain filling rate (Liu et al., 2019). Sucrose is primarily transported from source to sink tissues via sieve elements (SEs) (Xu et al., 2017). Overexpressed OsNAC23 gene enhances sugar transport to sink organs and lowers levels of sucrose, nonstructural carbohydrate, and temporary starch accumulation. In contrast, its absence has the opposite effect in mutants (Li et al., 2022). Shi et al. (2013) evaluated nitrogen (N) and nonstructural carbohydrate translocation into grains, impacting yield, grain-filling dynamics, and quality in N22 and Gharib (heat-sensitive) under HNT. Reduced grain yield in Gharib resulted from decreased N and nonstructural carbohydrate translocation post-flowering, affecting grain-filling rate, weight, and quality. Enhanced HNT tolerance in N22 correlated with increased HSPs and calcium-signaling proteins, along with efficient protein modification and repair mechanisms, particularly during early grain-filling. Accelerated grain-filling rate and improved proteomic protection, supported by enhanced assimilate translocation, conferred HNT tolerance in rice. Thus, dynamic proteome programming across key developmental stages guides future crop improvement efforts. The metabolomic analysis provided potential markers for breeding stress-tolerant rice germplasm (Lawas et al., 2019).

8 How does the rice plant respond to HTS at the molecular level?

A slight increase in average temperature may not cause severe damage to cells, tissues, and heat shock responses (HSR). Still, it may cause morphological changes, biorhythms, and immunity response (Shrestha et al., 2022). The cell wall is the first protective barrier in plants and responds first to HS, followed by changes in membrane fluidity and activation of channel receptors, which play an essential role in thermal sensing, cellular response, and calcium signal transduction. Increased ROS levels prompt the release of Ca²⁺ from different organelles, resulting in the sudden release of apoplastic Ca²⁺ and raising the amount of free Ca²⁺ in the cytoplasm, which performs an essential role in activating or repressing Ca²⁺/Cam-related kinases, phosphatases, and transcription factors (TFs) (Kan and Lin, 2021). The cell wall remodeling protein, pectin methylesterase (PME), can have its expression upregulated by HS, which increases the amount of demethylesterified pectin. PME activity and pectin demethylesterification are escalated by increased H₂O₂ content, which ultimately activates the downstream regulatory pathway of HS response (Wu et al., 2018; Xu et al., 2021). OsCNGC14 and OsCNGC16 responded to HS by triggering calcium signals, and both mutants (oscngc14 and oscngc16) exhibited impaired Ca²⁺ influx to the cytosol and decreased or abolished cytosolic Ca²⁺-mediated signal transduction in response to HS (Cui et al., 2020) (Figure 6). Although OsCNGC9 and OsCNGC13 channels are likewise Ca²⁺ permeable and involved in cytosolic Ca²⁺ mediated signaling cascades, additional research is essential to fully elucidate their roles in HS response (Wang et al., 2019; Xu et al., 2017). The OsHsfA2e gene has been observed to be associated with a CaMK engaged in the Ca2+/calmodulin-dependent signaling pathway, and its interaction with Hsp70 and Hsp90 is crucial in the response to HS (Harshada et al., 2021). Changes in membrane fluidity induced by heat may activate PM-bound phospholipases and kinases, leading to rapid increases in phosphatidic acid and phosphatidylinositol diphosphate (PIP2), which are vital in stress signal transduction (Niu and Xiang, 2018).

Figure 6

Figure 6. General molecular responses to HS in rice and other crop plants. The cell wall responds first to HS and plays a crucial role in thermal sensing. Ca²⁺ signal induction is the most rapid response. The calcium signaling pathway is crucial for activating, inhibiting, and regulating various Ca²⁺/calmodulin-dependent kinases (CaMKs), phosphatases, and numerous TFs. Hwi1 acts as potential receptor-like kinase that function upstream in the heat-response pathway. Cyclic nucleotide-gated channels OsCNGC14 and OsCNGC16 respond to HS by triggering Ca²⁺ signals. TT2 facilitates the heat-induced rise in cytosolic Ca²⁺ levels, which is then interpreted through Cam-SCT1/2 interactions, suppressing OsWR2 (Wax Synthesis Regulatory2) transcription. RGB1 (encodes a β-subunit of G protein) reduces electrolyte leakage and the accumulation of MDA and H₂O₂. RGA1 encodes the Gα subunit of the G protein and is involved in signal transduction. During HS, TT3.1 moves from the plasma membrane to the endosomes, where it recruits TT3.2 and facilitates its degradation through ubiquitination, thereby preventing the accumulation of mature TT3.2 in the chloroplast and protecting thylakoids. In response to HS, ZmCDPK7 (calcium-dependent protein kinase) translocates from the PM to the cytosol and phosphorylates sHSP17.4 (small HSP 17.4) and RBOH (a key producer of ROS), enhancing their expression levels. HS triggers translocation of OsNTL3 (a NAM, ATAF1/2, CUC1/2[NAC] transcription factor) from PM to the nucleus and activates transcription of downstream genes. ANN1 (annexin1), SNAC3, DST, and HTAS are pivotal in the scavenging of ROS by regulating the accumulation of key antioxidant enzymes like CATs (Catalases) and SOD (Superoxide dismutases) and APXs (Ascorbate peroxidases). The endoplasmic reticulum-localized TFs OsbZIP74 and ZmbZIP60 are spliced by IRE1 and are involved in the transcriptional activation of OsNTL3 and ZmHUG1(Heat Up-Regulated Gene 1). The PSII components D1, OsIF (encodes intermediate filament protein), PGL (encodes chlorophyllide a oxygenase 1), and HES1 (UGPase enzymatic activity) are essential for preserving chloroplast ultrastructure. During HS conditions, TSCD11 (seryl t-RNA synthetase) and Hsp70CP1 (chloroplast localized Hsp70) regulate the chloroplast development. HYR, a transcription factor and NSUN2 (RNA methyltransferase) ensure the transcripts of the photosynthetic gene, while HTS1(β-ketoacyl carrier protein reductase localized in thylakoid membrane) regulates lipid metabolic flux. OsTRXz–OsFLN2–OsFLN1 complex (encoding thioredoxin protein and fructokinase-like proteins), PGL, HTS1, HES1, and TSCD11 work together to maintain redox balance in the chloroplast. TT1 and HTAS remove unfolded proteins, while the RACK1A–eIF3h–AET1 (Receptor for Activated C Kinase 1A- eukaryotic Initiation Factor Subunit h- Adaptation to Environmental Temperature 1) complex, SLG1, and TOGR1 ensure protein translation under HS. In response to HS, the HSP-HSF complex dissociates, releasing HSFs. This release removes the NF-YA9/A10 (Nuclear Factor Y subunit A9/A10) - mediated inhibition of heat shock response (HSR) through HSF-driven accumulation of miR169s. GSA1 enhances thermotolerance in rice by regulating the flux of flavonoid glycosides and anthocyanins. In maize, GOLS2 and RAFS are transcriptionally activated by HSF, which enhances raffinose biosynthesis to modulate heat tolerance.

During HS, ER-localized OsbZIP74 translocates to the nucleus, initiating the expression of OsNTL3, followed by the transport of membrane-localized OsNTL3 to the nucleus to regulate the expression of OsZIP74. OsNTL3 encodes a NAC TF with a predicted C-terminal transmembrane domain (Liu et al., 2020; Lu et al., 2012). The repair of damaged PSII and the PGL gene (pale green leaf, encoding chlorophyllide and oxygenase 1) in the chloroplast requires the de novo synthesis of the D1 subunit (Kan and Lin, 2021); meanwhile, the OsTRXz-OsFLN1/2, encoding thioredoxin protein and fructokinase-like proteins, respectively, complex shields chloroplasts from heat-induced damage (Lv et al., 2017) (Figure 6) and the mitochondria-localized EG1 protects the organelle and maintains floral robustness (Zhang et al., 2016).

The most significant elements of the intricate transcriptional regulatory network of HSR in plants are heat shock TFs, or heat shock transcription factors (HSFs). They initiate a transcriptional cascade that activates genes encoding HSR-induced TFs, ROS-scavenging enzymes, metabolic enzymes, and HSPs (Gong et al., 2020). Among 25 HSFs in rice, HT stimulates the expression of 22 of these genes (Mittal et al., 2009; Xu et al., 2021). Among these HSFs, HSFA1s are considered ‘‘master regulators” in the transcriptional network (Ohama et al., 2017). When rice plants experience HS, alternative splicing induces the transcriptionally active form of OsHSFA2d, which increases the expression of genes like HSP17.7, HSP18.2, HSP21, HSP83.1, and HSP101 (Cheng et al., 2015). The isolated proteins OsHSP17.4 and OsHSP17.9A exhibit chaperone activity by effectively preventing the aggregation of proteins (Sarkar et al., 2019).

9 Genetics of rice thermotolerance: key QTLs and candidate genes

HS tolerance is often viewed as a quantitative trait, and despite considerable research over the past several years, the genetic basis of HS tolerance remains largely unclear. In recent years, advancements in molecular marker technology and increased research focus on rice heat tolerance have identified numerous heat-tolerant QTLs across 12 rice chromosomes. Identifying QTLs for rice thermotolerance has been hampered by several issues, such as inconsistent phenotyping, lack of genetic resources with established thermotolerance, trait complexity, and environmental factors. Notwithstanding these obstacles, rice QTL mapping for heat tolerance has advanced significantly.

9.1 QTLs identified for thermotolerance in rice

The detection of QTLs helped understand rice’s genetic mechanism, marker-assisted selection, and QTL cloning (Khan et al., 2019). Efforts for molecular mapping of rice thermotolerant QTLs have been carried out at booting, flowering, and grain filling to ripening stages (Buu et al., 2021; Liu et al., 2023). Most recently identified heat-tolerant QTLs pertain to these stages, with fewer associated with seedling stages (Table 2). A natural QTL, TT2 (THERMOTOLERANCE 2), encoding a Gγ subunit, reduces yield losses under HS at both vegetative and reproductive stages. It regulates G protein, wax metabolism, and Ca²⁺ signaling, potentially reducing the yield penalty under HS (Kan et al., 2022) (Table 2). Major QTL, TT1-2, was controlled by a single dominant gene. It was narrowed to a 26.0 kb region (Yan et al., 2021). The QTL qEMF3, identified from wild rice O. officinalis, holds a very promising role in shifting rice cultivars’ flower opening time to earlier in the morning (Hirabayashi et al., 2015). By creating a mapping population using a common wild rice as one of the parents, heat-tolerant QTLs qHTH5, qHTH10, and qHTB1–1 were detected. At the heading and flowering stage, QTL qHTH5 was located within 304.2 kb on chromosome 5 short arm (Hu et al., 2022). SNP markers were utilized for mapping two major QTLs, namely qHTSF1.1 and qHTSF4.1, on chromosomes 1 and 4, respectively by generating the population from a cross IR64/N22 for spikelet fertility variation (Ye et al., 2011). Two major QTLs for chalkiness, qPGC5 and qPGC6, were located by substitution mapping of single-segment substitution lines (SSSLs). qPGC5 was located in the 876.5 kb interval of chromosome 5, and qPGC6 was located in the 269.1 kb interval of chromosome 6 (Yang et al., 2022). Fengfeng et al. (2023) mapped three heat tolerant QTLs at seedling stage from O. longistaminata, including novel heat tolerant loci qTT4 and qTT5. Two QTLs for spikelet sterility with significant genetic effects, qSTIPSS9.1 and qSTIY5.1/qSSIY5.2, were located within genomic regions below 400 kbp (PS et al., 2017). Park et al. (2020) utilized the double-haploid line derived from Cheongcheong/Nagdong to investigate the QTL associated with booting stage thermotolerance (Table 2). Major QTLs qHAC4, qHAC8a, qHAC8b, and qHAC10 minimize the adverse impact of HT on amylose content by upregulating the splicing efficiency of the Wx gene (Zhang et al., 2014). Eleven QTLs associated with thermotolerance at anthesis were detected by Zhao et al. (2016), further confirming qPSL^ht4.1 across various temperature conditions. This locus has been consistently detected in multiple studies. Numerous QTLs contributing to heat tolerance at grain filling stage have also been mapped. For instance, the Appearance quality of brown rice 1 (Apq1) QTL has been localized to a 19.4-kb region, with the underlying gene identified as sucrose synthase 3 (Sus3) (Takehara et al., 2018).

Table 2

Table 2. Major QTLs governing stress tolerance in rice.

9.2 Candidate genes and molecular insights for heat tolerance

Two major candidate genes LOC_Os08g07010 and LOC_Os08g07440 identified within a major QTL qHTT8, on chromosome 8, controlling flowering stage thermotolerance (Chen et al., 2021). Gene LOC_Os09g38500 within the novel QTL qRSF9.2 region is associated with controlling relative spikelet fertility under HS (Hu et al., 2022). The qHTB1–1 QTL, controlling thermotolerance at the booting stage in rice, was fine-mapped to a 47.1 kb region containing eight candidate genes. Two positional candidate genes (LOC_Os01g53160 and LOC_Os01g53220) showed significant changes in expression levels under HS (Cao et al., 2020). OsHTAS codes for a ubiquitin ligase found in both the nucleus and cytoplasm. It reacts to various stresses and shows robust activation in response to externally applied ABA. OsHTAS regulates the accumulation of H₂O₂ in shoots, affects the opening of stomata on rice leaves, and enhances ABA biosynthesis (Liu et al., 2016) (Table 3). Candidate genes LOC_Os04g52830 and LOC_Os04g52870 located within the locus qHTT4.2 enhance seed setting rate under HS (Pan et al., 2023). Das et al. (2024) carried out genome-wide association mapping and found three significant QTLs and three promising putative candidate genes regulating the Photosystem II (PSII) complex impairment during HS. Common wild rice (O. rufipogon Griff.) is a valuable source of germplasm that can be used to improve rice. The genetic population created with the common wild rice (O. rufipogon Griff.) as a parent is a robust QTL mapping population for HS tolerance. In addition, common wild rice is a significant source of high-temperature tolerant rice germplasm (Ishimaru et al., 2010; Xiao et al., 1996).

Table 3

Table 3. Stage specific genes involved in HS tolerance across growth stages in rice.

Knockout mutants of OsRboh (NADPH oxidase) showed upregulation of heat responsive genes in seedlings of Nipponbare (Liu et al., 2025). At seedling stage OsIAA7 increases thermotolerance by positively affecting malondialdehyde, catalase, and chlorophyll A levels. It decreases H₂O₂ levels and prevents cell death. OsARF6 (auxin response factor gene) negatively modulates heat tolerance by influencing the expression levels of OsTT1 and OsTT3.1 (Qiu et al., 2025). Two locus namely qDW7 (dry weight) and qFW6 (fresh weight) showed association with rice response to HT. Localized genes such as LOC_Os06g10790, LOC_Os07g30330, and LOC_Os03g59040 were found responsive to HS at seedling stage (Wei et al., 2021). Expression level of genes (ethylene insensitive 2, ethylene insensitive-like1, and ethylene insensitive-like 2) involved in ethylene signaling, is increased under HS in rice seedlings (Wu and Yang, 2019).

In seeds developed during HS, the promoter regions of starch biosynthesis genes OsAGPS2b (ADP-GLc pyrophosphorylase subunit 2b), OsGBSSI (granule-bound starch synthase), and OsSuSy2 (sucrose synthase 2) showed marked hypomethylation. In contrast the promoters of α-amylase genes OsAmy1A and OsAmy3D were significantly hypermethylated (Suriyasak et al., 2025). A promotor variant of WCR1 (White-Core Rate 1) increases OsDOF17 binding, leading to higher WCR1 expression, which in turn reduces chalkiness and enhances grain quality (Wu et al., 2022). HS upregulated the expression of genes associated with NADPH oxidases (OsRbohB, OsRbohD, OsRbohF, and OsRbohl) and GA biosynthesis (OsGA30x1 and OsGA20ox1) during grain filling. The GSA1 (Grain Size and Abiotic Stress tolerance 1) locus, which encodes a UDP-glucosyltransferase, controls grain size by influencing cell proliferation and cell expansion (Dong et al., 2020). Genes ONAC127 and ONAC129, encoding TFs are expressed in pericarp modulating sugar transport during HTS at grain development stage (Ren et al., 2021) (Table 3).

10 Heat stress-induced epigenetic modification in rice

In addition to morphological and molecular adaptations, plants also adapt through epigenetic mechanisms, which involve DNA methylation, chromatin modeling, histone modification, sRNAs, and lncRNAs for surviving under adverse environmental conditions by altering the gene expression pattern and/or epigenetic memory (Crisp et al., 2016; Kinoshita and Seki, 2014; McCormick, 2018). Epigenetics has a potential role in increasing thermotolerance, as demonstrated by the growing acceptance of its application in HS resilience breeding (Gardiner et al., 2015; Yadav et al., 2022).

During rice seed development, moderate HS causes the demethylation of the OsFIE1 (O. sativa Fertilization-Independent Endosperm1) locus and represses CMT3 (CHROMOMETHYLASE3) expression, leading to correlated increase in OsFIE1 transcript levels. This increase is associated with a decrease in repressive H3K9me2 mark on OsFIE1. Enhanced OsFIE1 activity (as part of Polycomb Repressive Complex 2) then deposits the repressive H3K27me3 (Histone H3 lysine 27 trimethylation, an epigenetic modification) mark on target genes like MADS82 and MADS87, (MADS box TFs), which ultimately causes precocious cellularization and reduced seed size (Folsom et al., 2014). The indica rice variety 93–11 shows more dynamic changes in chromatin accessibility and gene expression under HS than the japonica variety Nipponbare, aligning with the greater heat tolerance observed in 93–11 relative to Nipponbare (Liang et al., 2021). Genome-wide survey of histone H3 lysine4 tri-methylation (H3K4me3) under drought conditions revealed differential methylation of 4837 genes, out of which 3927 showed increased expression while 910 showed decreased total transcript. HS during grain filling significantly increased DNA methylation of promotors in ABA metabolism related genes (OsNCED2, OsNCED3, and OsNCED5), ABA catabolism genes (OsABA8′OH1, OsABA8′OH2, and OsABA8′OH3), and α-amylase genes (OsAmy1C, OsAmy3B, and OsAmy3) causing delayed germination of heat stressed seeds. Predicted CpG islands were found in these genes except in genes OsNCED2, OsABA8′OH2, or OsAmy1A. Presence of CpG islands and hyper-methylation in α-amylase promotors cause HS induced transcriptional regulation during seed imbibition (Suriyasak et al., 2020). Under HS, rice roots and shoots displayed varied expression levels of miR160 and miR169, indicating that heat regulates target genes differently in these two distinct tissues (Sailaja et al., 2014). Elevated temperatures during early seed development prompted genome-wide alterations, including the reactivation of transposable elements by a decrease in DNA methylation in noncoding regions (Chen et al., 2016). Rice lines overexpressing tae-miR159 and Arabidopsis myb33myb65 double mutants exhibit increased sensitivity to HS compared to their wild-type counterparts. This suggests that the down-regulation of miR159 and the subsequent up-regulation of its target genes following HS may be involved in a HS-related signaling pathway, contributing to HS tolerance (Wang et al., 2012). Li et al. (2023) applied methylation-sensitive amplification polymorphism31 (MSAP) to investigate the DNA methylation responses in rice spikelets at the anthesis stage under control and HT conditions. DNA methylation level significantly increased in the susceptible rice group and decreased in the tolerant rice group under HT treatment. Genes in N22 leaves with hypomethylation that exhibit up-regulated expression under direct seeded rice circumstances, implying that epigenetic alteration plays a significant role in N22’s adaptive plasticity. The ability of N22 to adapt to unfavorable climatic conditions is attributed to variations in chromatin architecture and post-translational modification of proteins, including histone modifications (Seem et al., 2024). Liu et al. (2017) mapped four QTLs (qHT-3, qHT-6, qHT-8, and qHT-12) for thermotolerance during flowering. Their detailed analysis revealed that these QTLs contained miRNA targets associated with ABA-responsive genes. Moreover, they discovered that the suppressor of the G2 allele of skp1 (SGT1), a direct target of miRNA166e, was located within the qHT-8 locus. Studies of the mechanism and nature of these various epigenetic modifications can provide valuable insights into the different genes and the specific regions in them responsible for adapting to HTS, leading to the development of a better understanding of the pathway to be targeted for thermotolerance breeding.

11 Breeding to improve thermotolerance in rice

Among the various strategies aimed at alleviating the impact of HS on rice, breeding emerges as a fundamental approach. Development of heat-tolerant rice cultivars through thermotolerance targeted breeding programs presents a cost effective, long-lasting and sustainable solution. Relative to other abiotic stresses like drought and salinity, breeding endeavors for heat-tolerant rice varieties have attracted less research and focus. There is inadequate data concerning HS resistant rice cultivars and the genetic mechanisms underlying their tolerance. Moreover, the identified genetic resources and QTLs have not been comprehensively employed in breeding programs to develop thermotolerant rice varieties (Stephen et al., 2022).

11.1 Conventional breeding

Conventional breeding typically relies upon extensive phenotypic characterization, selecting phenotypes related to thermotolerance, and is conducted in regions with climates similar to where the crop will be cultivated (Driedonks et al., 2016). Assessing thermotolerance levels precisely, selecting superior breeding lines, and effectively transferring traits related to heat tolerance into specific cultivars with favorable agronomic traits are crucial aspects of conventional breeding. The straightforward traits, such as pollen fertility, seed setting rate, spikelet fertility, grain chalkiness, etc., serve as reliable indicators in conventional breeding for heat tolerance. Using these indicators, a large number of heat-tolerant rice genotypes have been identified and some has been utilized as donors to develop breeding lines, including N22 (Kilasi et al., 2018; Ye et al., 2015), Giza178 (Abdelaal et al., 2021), IR2061 (Cheng et al., 2012), and (Zhao et al., 2016) (Table 2). Some heat-tolerant hybrid rice varieties, such as Guodao 6, maintain seed-setting stability under HS conditions due to their adaptability to heat avoidance, characterized by floral traits like shortened flowering phases (Tao et al., 2008). N22 is frequently employed as a control in studies focusing on heat tolerance. Giza178, originating from a japonica-indica cross and hailing from Egypt, demonstrates notable heat tolerance (Tenorio et al., 2013). Interspecific rice hybrid NERICA-L-44 has been characterized as a heat-tolerant variety (Bahuguna et al., 2015). Using ²⁰Gy proton, three heat-tolerant mutant lines 8852, 8552, and LP-12 were developed utilizing varieties J-104 and A-82 through a mutation breeding approach (Gonzalez et al., 2021).

11.2 Marker-assisted selection for thermotolerance in rice

Improving yield under stress conditions through direct selection is constrained by the low heritability and intricate nature of the QTLs that govern them. Marker-assisted selection (MAS) has proven effective in precisely transferring genes from wild germplasm while minimizing linkage drag, as it can integrate modern classical genetics, bioinformatics, and conventional biotechnology effectively. Techniques such as association mapping and bi-parental mapping can elucidate the connection between phenotypic variation and genetic polymorphism, facilitating the mapping of relevant genomic regions (Visakh et al., 2024). After mapping QTL regions associated with thermotolerance, these regions can be incorporated into superior genetic backgrounds employing MAS in breeding programs. Utilizing marker-assisted backcrossing (MABC), a few lines have been developed for HS tolerance. For example, Lang et al. (2015) applied MABC using six markers to enhance heat tolerance in rice varieties in Vietnam. They crossed heat-tolerant germplasms (N22 and Dular) with five high-yielding indica cultivars, resulting in the development of four heat-tolerant lines by the BC₄F₂ generation. Ye et al. (2022) introgressed two QTLs, namely qEMF3 and qHTSF4.1, for early morning flowering and thermotolerance, respectively, into the background of IR64, and a QTL pyramiding line IR64+qHTSF4.1+qEMF3 was developed using MAS. In another effort in marker-assisted pedigree breeding by Withanawasam et al. (2022), they developed PL457 and PL130 with heat-tolerant QTLs qSSP_F10 and qHT6. These lines showed 85.02% and 61.55% yield advantages, respectively over IR64. Similarly, Vivitha et al. (2017) utilized MABC to introduce QTLs (qHTSF1.1 and qHTSF4.1) into Improved White Ponni lines, effectively enhancing thermotolerance. (Figure 7A).

Figure 7

Figure 7. Strategies to develop thermotolerance in rice. (A) Breeding approaches: Backcross (BC), marker-assisted selection (MAS), double haploid (DH), recombinant inbred lines (RILs), near-isogenic lines (NILs), and chromosome segments substitution lines (CSSLs) breeding methods are used extensively to develop varieties with desirable traits, including thermotolerance. Extensive phenotyping, characterization, and selection of germplasm for heat tolerance, assessing the thermotolerance level, and transferring HS tolerance into agronomically superior cultivars are the significant components of breeding for heat tolerance. (B) Omics Approaches: The incorporation of different omics technologies could broaden our understanding of molecular and biochemical insights into the complex interplay among genes, pre- and post-translational modifications, including DNA methylation, expressed proteins, and metabolites. These technologies help in the identification of heat-responsive QTLs, genes, proteins (HSPs, chaperones, dehydrins, LEA, etc.), and metabolites (prolines, sugars, spermidine, betaine, phenolic compounds, amino acids, and lipid-derived metabolites such as jasmonic acid). Hormonomics is a key tool for deep physiological phenotyping and could provide insights into cellular signaling, facilitate gene discovery, and allele mining. It is crucial to understand the microbiome and its interaction with the host under HS conditions. Deciphering whole genomes of microorganisms, i.e., microbiomics, is essential to uncovering the complex interaction between plants and microbes under stress conditions. (C) Genome editing (GE) approaches: GE is one of the most promising approaches to understanding and enhancing HS tolerance. Four significant steps could be taken to utilize these techniques, (I) Identifying HS-responsive genes, (II) Targeted gene editing, i.e., to protect photosynthesis (RuBisCo, RCA, D1, IF, HYR, and TT3.1–TT3.2), improving stomatal closure and cuticle deposition to prevent water loss (MDHAR4, HTAS, DST, and the TT2-SCT1-WR2 regulatory pathways), optimizing the heading/flowering stage (pathways MADS51/qHd1–Ehd1–RFT1/Hd3a and Ghd7–Ehd1–RFT1/Hd3a or specific genes such as EG1 and DFOT1), enhancing fertility (through increased osmoproctants, balanced ROS homeostasis, stress responsive metabolites) and maintaining normal endosperm development under HS to reduce grain chalkiness and enhance grain weight and quality (ONAC127/129, Fie1, MADS87, MADS57-Wx, DOF17–WCR1–MT2b (Metallothionein 2b), bZIP60, bZIP58–Amy1C/3A/3D, TT1, Hsp70cp-2, and ZmPGD3 homologs) through CRISPR-Cas9, ZFNs, TALENs etc. (III) Incorporation of edited gene into the plant cells, and (IV) Confirmation of genome editing in heat tolerant rice plant. (D) Data-driven approaches: Precise, accurate, reliable, and reproducible phenotyping is needed to identify the most important traits and promising donors with superior characteristics for use in HS tolerance breeding programs in rice. Mobile and stationary platforms are two primary categories in high-throughput phenotyping platforms. Mobile platforms include field-based platforms (satellite imagery, UAV imaging, and ground-based imaging through PhenoMobile) and laboratory-based platforms (MVS-Pheno). Stationary platforms include facilities such as the Nanaji Deshmukh Plant Phenomics Centre (NDPPC), New Delhi. Imaging techniques are an essential part of high-throughput phenotyping (HTP). Magnetic Resonance Imaging (MRI), X-ray/CT (X-ray Computed Tomography), RGB (Visible light), Infrared thermal (IR) imaging, fluorescence imaging, and LiDAR (Light Detection and Ranging). These HTP facilities can transform HS phenotyping and screening with high spatial and temporal resolution. By combining machine learning (ML) and deep learning (DL) in HTP, we may be able to use deep neural networks to evaluate big data sets and images with more precision and accuracy.

11.3 Genome editing and transgenic approach

Genome editing technologies in crop plants are advancing rapidly, allowing for targeted mutations with exceptional specificity and accuracy. Unlike genetic engineering, genome editing doesn’t involve the integration of foreign DNA into plants, making the final product indistinguishable from the parent plants without changing the overall stability of the genome. Genome editing tools such as zinc finger nucleases (ZFN), meganucleases, transcription activator-like effector nucleases (TALEN), and CRISPR/Cas system offer promising avenues for creating improved plant varieties by adding desirable traits or removing undesirable ones (Noman et al., 2016). Among these, CRISPR system stands out for its precision in modifying DNA at specific locations. It is being extensively utilized for the rapid, easy, and efficient alteration of genes to enhance HS in rice and to understand the function of specific genes by creating gene knockout mutants (Biswal et al., 2019; Tingting et al., 2023).

Qu et al. (2021) co-overexpressed RuBisCo and RuBisCo activase in rice plants and analyzed the photosynthesis and biomass at 25°C and 40°C. Plants showed a higher CO₂ assimilation rate at 40°C, which resulted in 26% higher dry weight than the wild types. To stimulate β-glucuronidase (GUS) gene expression in rice, promoters of the three highly heat-inducible genes OsHsfB2cp (a HSF), PM19p (a stress responsive promoter), and Hsp90p were employed. The transgenic rice panicles and flag leaves validated high heat-induced GUS activities, mild drought-induced activities through GUS gene expression and histochemical staining. While OsHsfB2cp and PM19p showed significantly higher activities in panicles under HS, the three promoters displayed comparable high activity levels in rice leaves (Rerksiri et al., 2013). OsDHSRP1 (an E3-ubiquitin ligase) regulates plant abiotic stress tolerance via the Ub/26S proteasome system, and its transcripts are highly expressed in heat and drought conditions. The Arabidopsis showed hypersensitivity to heat by reducing its germination rates and root length when OsDHSRP1 was overexpressed. It suggests that OsDHSRP1 E3 ligase acts as a negative regulator, and the degradation of its substrate proteins via ubiquitination has an essential role in the modulation of HS response via an ABA-independent pathway (Kim et al., 2020). When ERECTA (receptor-like kinase)-overexpressing plants were subjected to HT (42°C Day/35°C night) for 10 days at the reproductive stage, most leaves and tillers of retained their green color and survived as compared control lines. Additionally, the seed-setting rate of ERECTA-overexpressing plants was 55%–70% higher than the control line following HTS (Shen et al., 2015) (Table 4). When a zinc finger protein gene, OsZFP350, was upregulated, transgenic rice plants showed enhanced primary root length, adventitious and lateral roots, and significantly increased germination rate (Kang et al., 2019).

Table 4

Table 4. HS-tolerant transgenic rice: Genes and their functions.

Silencing of CI-sHsps (Class I small HSPs) by RNAi negatively affected rice seedlings’ seed germination process and HS response. The seed length was reduced, the seed germination process was delayed, and seed thermotolerance was negatively affected compared to non-transgenic seeds (Sarkar et al., 2019). Eliminating ABA receptor (PYL1/4/6) encoding genes via CRISPR/Cas9 has conferred substantial thermotolerance in rice (Miao et al., 2018). Rice knockout mutants of OsHSP60-3B demonstrated normal fertility under favorable temperatures but experienced fertility reduction with rising temperatures. Pollen from mutant oshsp60-3b shows decreased starch accumulation and viability, while anthers produce enhanced levels of ROS, resulting in cell death. Further exploration of this gene and its implications for HS response is warranted (Lin et al., 2023).

Genome editing provides a fast approach for developing thermotolerant rice varieties compared to conventional methods by precisely targeting specific genes, thus enabling fine-tuning tolerance. If these advanced approaches are suitably amalgamated with techniques such as speed breeding and genome-wide association studies (GWAS), it will revolutionize the efforts for developing HS-resilient rice varieties (Figure 7C).

11.4 Omics approaches for developing heat-tolerant rice

Large-scale, high-throughput approaches such as genomics, transcriptomics, proteomics, metabolomics, epigenomics, hormonomics, ionomics, and phenomics are central to omics technology. Omics in rice research has deepened the understanding of complex molecular responses, differential gene expression, regulatory pathways, comprehensive genome-wide insights into gene structures, their functions, interconnected regulatory networks, metabolic and biochemical processes they participate in, thereby helping to elucidate how gene networks interact with complex stress resilient traits with potential applications in crop improvement for thermotolerance (Iqbal et al., 2021; Pandian et al., 2020; Singh et al., 2024; Varshney et al., 2018; Zhang et al., 2019).

Proteomic studies have shown stage specific patterns of differentially expressed proteins at various developmental stages under HTS. These proteins are primarily associated with biosynthesis, energy and metabolic processes, redox balance, photosynthesis, and signaling pathways, and contribute to short term protective response that enhance thermotolerance (Guo et al., 2024). In various extensive studies, heat-responsive QTLs and genes have been identified and cloned, such as by Huang et al. (2023) and Nguyen et al. (2022). Li et al. (2023) identified a major locus qHT7 through GWAS and further transcriptome analysis revealed candidate gene LOC_Os07g48710 (OsVQ30, a transcriptional regulatory factor) within this locus. Liu et al. (2021) studied regulatory effects of HT on grain development and material accumulation pathways. Proteomic findings revealed a total of 840 differentially expressed proteins during the grain filling process. Proteins such as PPROL 14E, PSB28, granule-bound starch synthase I were upregulated.

Transcriptome of a rice hybrid showed upregulation of genes involved in responses to stimuli, cell communication, and metabolic and TF activities while down regulated genes were enriched in photosynthesis and signal transduction (Wang et al., 2020). Heat-tolerant genotype, SDWG005, maintained a steady state balance of metabolic processes, associated with reprogramming cellular activities and had a critical role in preserving heat tolerance (Cai et al., 2020) (Table 5). HS changes phosphorylation dynamically and alters its pattern in enzymes related to starch biosynthesis (Pang et al., 2021). Roots of heat tolerant cultivar mainly activated pathways involving phenylalanine/phenylpropanoid, aromatic amino acid, lysine degradation, branched chain amino acids, glycerophospholipids, and alkaloids while heat susceptible cultivar emphasized nitrogen-related and antioxidant pathways under HTS in a metabolomic study by Ogawa et al. (2025) (Table 5). Targeted metabolomic studies are essential to elucidate inter-organ communication and hormonal crosstalk in rice during HTS (Prerostova et al., 2022).

Table 5

Table 5. Recent studies on HS tolerance in rice using different omics technologies.

HS alters various aspects of chromosomes organization, including shifts between A and B compartments, enlarged topologically associated domains, and reduced short range chromatin interactions. These structural modifications correlate with the changes in chromatin accessibility and gene expression (Liang et al., 2021). HS significantly alters splicing patterns in rice. Two notable candidate genes, LOC_Os03g16460 (encoding an uncharacterized protein) and LOC_Os05g07050 (encodes pre-mRNA splicing factor 8) were linked to alternate splicing and intron retention (IR) type of alternate splicing being more common under HS (Yang et al., 2022). A rice specific DOF transcription factor, OsDOF27 (an intrinsically disordered protein) is highly regulated under HS. Its promoter region in enriched with abiotic stress and phytohormone-responsive cis-elements (Gandass et al., 2022). Ubiquitination, a major post-translational modification, contributes critically to thermotolerance. Ying et al. (2023) identified 488 ubiquitination sites in 246 proteins in rice endosperm during HTS in a ubiquitomic study. Ubiquitination likely contributes to decreased abundance of starch synthesis enzymes, explaining reduced amylose content under HS.

Studies using omics technologies in rice have revealed significant genetic and molecular factors that help to understand the intricate mechanism of HS tolerance (Table 5). Developments in computational biology and bioinformatics have significantly improved the analysis and interpretation of proteomic data. Machine learning tools for network analysis and algorithms determine the critical regulatory proteins and their interaction, shedding light on the intricate regulatory networks. These holistic approaches has resulted in identifying possible protein targets for breeding and genetic engineering to create rice cultivars that can withstand HT (Jhan et al., 2023; Singh et al., 2024) (Figure 7B).

11.5 Harnessing plant-microbe interactions to improve thermotolerance in rice

Rice plants’ microbiome consists of diverse microorganisms (bacteria, archaea, fungi, and viruses) present both inside and outside of their tissues, in the endosphere and ectosphere, respectively (Singh et al., 2022; Aswini, 2023; Zhao et al., 2024). Beneficial micro-organisms can help to reprogram the plant epitranscriptome to enhance thermotolerance (Shekhawat et al., 2022). Waqas et al. (2015) reported that the endophytic fungus Paecilomyces formosus LWL1 produces phytohormones and organic acids, which enhanced the HS tolerance of a japonica variety, Dongjin. Bacillus amyloliquefaciens NBRI-SN13 (SN13) ameliorated various abiotic stresses, including HS in a rice variety Saryu-52 (Tiwari et al., 2017). In both the rhizosphere and endosphere, a significant decline in microbial abundance has been observed in the IR64 (temperature sensitive) cultivar compared to the Huanghazou (temperature resistant) cultivar (Munir et al., 2023). Rice plants inoculated with Brevibacterium linens RS16, which produces 1-aminocyclopropane-1-carboxylate (ACC) deaminase, had enhanced tolerance against UV-b radiation, and 40°C of temperature because of decreased ethylene emissions, increased plant biomass and photosynthesis, and restricted DNA damage (Choi et al., 2022).

Microorganisms enhance HS tolerance by the production of antioxidants, plant growth hormones, bioactive compounds, detoxifying harmful compounds, sequestration of ROS and other free radicals, and releasing protective substances to withstand desiccation (Singh et al., 2023; Hosseiniyan Khatibi et al., 2024). So, understanding and deciphering the mechanics and interplay of the phyto-microbiome in rice with HS tolerance using the potential of multi-omics, e.g., metagenomics and meta-transcriptomics approaches, holds great promise to uncover the complex microbial network involved in stress signaling and development of stress tolerance (Figure 7B).

12 Integrating high-throughput phenotyping and artificial intelligence for HS tolerance breeding

The complexities imposed by HTS at the cellular level are further compounded at the physiological level by interactions between phytohormones and gene-regulatory proteins. This complexity is further heightened with interactions between different temperatures (e.g., high day and night temperatures) and other abiotic stresses, such as drought. Thus, accurately and precisely measuring traits impacted by HTS necessitates careful observation and interpretation, often in real-time. High throughput phenotyping (HTP), which uses imaging techniques, including fluorescence, visible range, near-infrared, laser, thermal, and hyperspectral imaging, has enabled measuring traits with extreme precision and at a high temporal and spatial scales in a non-destructive manner on a large scale to characterize crop response to HT (Bahuguna and Jagadish, 2015; Kundu et al., 2024). In addition to that, conventional plant phenotyping is time-consuming, labor-intensive, and expensive. HTP can be further improved by integrating artificial intelligence (AI) techniques for collecting reliable data to identify promising traits and genotypes for faster genetic gain and increased heritability. Deep phenotyping enables observing changes in tissue composition (e.g., proteins, lipids, carbohydrates, and metabolites) in vivo (Kundu et al., 2024). High frequency, high resolution imaging using a unmanned aerial vehicle (UAV) quantified dynamic drought responses of a rice population under field conditions using deep convolutional neural networks (DCNNs) together with canopy heights models. Traits such as UAV-based leaf rolling score, plant water content, and drought resistant index by UAV were measured. Genome wide association analysis identified 111 significant loci associated with three dynamic traits (Jiang et al., 2021). Convolutional neural networks (CNNs) and Gradient-weighted Class Activation Mapping (Grad-CAM) based approach was used to detect chalkiness in rice grains (Wang et al., 2022). Payman et al. (2018) used computer vision system to assess rice appearance qualities such as whiteness and chalkiness. Thermal stressed-induced spikelet sterility was investigated using a crop-model-assisted GWAS approach. Parameters provided RIDEV model was able to give more and strong QTLs then traits taken by observation (Dingkuhn et al., 2017). A high-throughput chlorophyll fluorescence platform enables rapid screening of photosynthetic heat tolerance in rice, revealing highly heritable PSII heat tolerant traits, substantial genetic variation across diverse germplasm, and 133 candidate genes underlying PSII thermotolerance through GWAS (Robson et al., 2023).

Combining HTP, AI, deep learning, and integrated data networks can help developing large repositories of genotyping data, producing big and accurate phenotypic data sets (Gill et al., 2022; Khatibi and Ali, 2024), accelerating the pace of developing stress resilience and discovering new traits and genes for HS tolerance as shown in Figure 7D.

13 Conclusion

Increasing efforts to develop heat-tolerant varieties become critical for food security amid climate change (Kan et al., 2023). Plenty of information is available about the physiological and metabolic traits that govern heat tolerance, but the underlying complex mechanisms are still unknown. Construction of high-density QTL maps for heat tolerance has provided a strong foundation. However, effective utilization of genetic resources, which have already been identified in breeding programs, remains a challenge. Approaches such as transgenics and advanced molecular breeding play a critical role in determining rice’s responsible genes and complex pathways governing HS tolerance. Genome editing techniques such as CRISPR/Cas9 and RNAi could be promising techniques. Advances in sequencing and high-throughput omics technologies have led to the generation of a vast array of omics data. Genomic regions governing heat tolerance can be identified by genomics and genetic mapping. Integrative studies of different disciplines such as structural genomics, proteomics, transcriptomics, and metabolomics with rice physiology and breeding would accelerate the efforts to identify key proteins, novel marker genes, and metabolic pathways to elucidate molecular mechanisms of thermotolerance and develop heat-tolerant rice varieties. Under stress conditions, various genes are expressed differentially, which is regulated by several processes, including N-tail modifications, histone variants, and DNA methylation. Comprehensive research of epigenetic regulatory mechanisms under HTS, particularly reproductive and grain-filling stages, needs to be extended and investigated in field conditions, as rice may encounter drought along with HS (Begcy and Dresselhaus, 2018). Incorporation of various HTP technologies integrated with AI tools assists in precise, detailed, deep, and large-scale phenotyping, considering the complexities imposed by HS. Microbial communities show greater resilience towards changes in the environment. Multi-omics approaches could play a key role in deciphering rice’s HS tolerance mechanism governed by plant and microbiome interactions. This integration of different methods in MAS and genomic selection will increase efficiency and accuracy in enhancing heat tolerance through breeding programs.

Author contributions

NK: Formal analysis, Writing – original draft, Visualization, Conceptualization, Validation, Writing – review & editing, Methodology, Investigation. SH: Validation, Writing – review & editing, Investigation, Project administration, Visualization, Formal analysis, Supervision, Writing – original draft, Methodology, Conceptualization. DS: Validation, Supervision, Methodology, Writing – review & editing. FA: Validation, Methodology, Writing – review & editing. GK: Methodology, Writing – review & editing, Validation. JA: Funding acquisition, Project administration, Visualization, Resources, Validation, Supervision, Conceptualization, Methodology, Investigation, Writing – review & editing, Writing – original draft.

Funding

The author(s) declared that financial support was received for this work and/or its publication. This publication was funded by the Ministry of Agriculture and Farmers Welfare, Government of Madhya Pradesh, India and the International Rice Research Institute (IRRI)-Hybrid Rice Development Consortium (HRDC).

Acknowledgments

The authors would like to thank everyone at IRRI who participated in the manuscript’s internal review. Their insightful recommendations have helped to make it more improved.

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Generative AI statement

The author(s) declared that Generative AI was not used in the creation of this manuscript.

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References

Abdelaal, K., Mazrou, Y., Mohamed, A., Ghazy, M., Barakat, M., Hafez, Y., et al. (2021). The different responses of rice genotypes to heat stress associated with morphological, chlorophyll and yield characteristics. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 49, 12550–12550. doi: 10.15835/nbha49412550

Crossref Full Text | Google Scholar

Ahmad, S., Tabassum, J., Sheng, Z., Lv, Y., Chen, W., Zeb, A., et al. (2024). Loss-of-function of PGL10 impairs photosynthesis and tolerance to high-temperature stress in rice. Physiologia Plantarum 176, e14369. doi: 10.1111/ppl.14369

PubMed Abstract | Crossref Full Text | Google Scholar

Akhter, D., Qin, R., Nath, U. K., Eshag, J., Jin, X., and Shi, C. (2019). A rice gene, osPL, encoding a MYB family transcription factor confers anthocyanin synthesis, heat stress response and hormonal signaling. Gene 699, 62–72. doi: 10.1016/j.gene.2019.03.013

PubMed Abstract | Crossref Full Text | Google Scholar

Ali, Z., Merrium, S., Habib-Ur-Rahman, M., Hakeem, S., Saddique, M. A. B., and Sher, M. A. (2022). “Wetting mechanism and morphological adaptation; leaf rolling enhancing atmospheric water acquisition in wheat crop-a review. Environ. Sci. pollut. Res. Int. 29, 30967–30855. doi: 10.1007/s11356-022-18846-3

PubMed Abstract | Crossref Full Text | Google Scholar

Ambavaram, M. M. R., Basu, S., Krishnan, A., Ramegowda, V., Batlang, U., Rahman, L., et al. (2014). Coordinated regulation of photosynthesis in rice increases yield and tolerance to environmental stress. Nat. Commun. 5, 5302. doi: 10.1038/ncomms6302

PubMed Abstract | Crossref Full Text | Google Scholar

Arshad, M. S., Farooq, M., Asch, F., Krishna, J. S.V., Prasad, P.V.V., and Siddique, K. H.M. (2017). Thermal stress impacts reproductive development and grain yield in rice. Plant Physiol. Biochemistry: PPB 115, 57–72. doi: 10.1016/j.plaphy.2017.03.011

PubMed Abstract | Crossref Full Text | Google Scholar

Aswini, K. (2023). Microbiome mediated management of heat stress in wheat (Triticum aestivum L.) (New Delhi (India): Division of Microbiology, ICAR-Indian Agricultural Research Institute).

Google Scholar

Bahuguna, R. N. and Jagadish, K. S.V. (2015). Temperature regulation of plant phenological development. Environ. Exp. Bot. 111, 83–90. doi: 10.1016/j.envexpbot.2014.10.007

Crossref Full Text | Google Scholar

Bahuguna, R. N., Jha, J., Pal, M., Shah, D., Lawas, L. M., Khetarpal, S., et al. (2015). Physiological and biochemical characterization of NERICA-L-44: A novel source of heat tolerance at the vegetative and reproductive stages in rice. Physiologia Plantarum 154, 543–559. doi: 10.1111/ppl.12299

PubMed Abstract | Crossref Full Text | Google Scholar

Bahuguna, R., Solis, C. A., Shi, W., and Jagadish, K. (2016). Post-flowering night respiration and altered sink activity account for high night temperature-induced grain yield and quality loss in rice (Oryza sativa). Physiologia Plantarum 159, 59–73. doi: 10.1111/ppl.12485

PubMed Abstract | Crossref Full Text | Google Scholar

Basak, J, Ali, M., Islam, M., and Alam, J. (2009). Assessment of the effect of climate change on boro rice production in Bangladesh using DSSAT model. Int. Center 38 (9), 95–108.

Google Scholar

Baxter, A., Mittler, R., and Suzuki, N. (2014). ROS as key players in plant stress signalling. J. Exp. Bot. 65, 1229–1405. doi: 10.1093/jxb/ert375

PubMed Abstract | Crossref Full Text | Google Scholar

Begcy, K. and Dresselhaus, T. (2018). “Epigenetic responses to abiotic stresses during reproductive development in cereals. Plant Reprod. 31, 343–555. doi: 10.1007/s00497-018-0343-4

PubMed Abstract | Crossref Full Text | Google Scholar

Begcy, K., Sandhu, J., and Walia, H. (2018). Transient heat stress during early seed development primes germination and seedling establishment in rice. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01768

PubMed Abstract | Crossref Full Text | Google Scholar

Berkeley Earth Data overview. Available online at: https://berkeleyearth.org/data/ (Accessed December 12, 2024).

Google Scholar

Biswal, A. K., Mangrauthia, S. K., Reddy, M.R., and Yugandhar, P. (2019). CRISPR mediated genome engineering to develop climate smart rice: challenges and opportunities. Semin. Cell Dev. Biol. 96, 100–106. doi: 10.1016/j.semcdb.2019.04.005

PubMed Abstract | Crossref Full Text | Google Scholar

Biswas, S., Islam, M. N., Sarker, S., Tuteja, N., and Seraj, Z. I. (2019). Overexpression of heterotrimeric G protein beta subunit gene (OsRGB1) confers both heat and salinity stress tolerance in rice. Plant Physiol. Biochem. 144, 334–344. doi: 10.1016/j.plaphy.2019.10.005

PubMed Abstract | Crossref Full Text | Google Scholar

Bita, C. E. and Gerats, T. (2013). Plant tolerance to high temperature in a changing environment: scientific fundamentals and production of heat stress-tolerant crops. Front. Plant Sci. 4. doi: 10.3389/fpls.2013.00273

PubMed Abstract | Crossref Full Text | Google Scholar

Brunel-Muguet, S., D'Hooghe, P., Bataillé, M. P., Larré, C., Kim, T. H., Trouverie, J., et al. (2015). Heat stress during seed filling interferes with sulfur restriction on grain composition and seed germination in oilseed rape (Brassica napus L.). Front. Plant Sci. 6. doi: 10.3389/fpls.2015.00213

PubMed Abstract | Crossref Full Text | Google Scholar

Buu, B. C., Chan, C. Y., and Lang, N. T. (2021). “Molecular breeding for improving heat stress tolerance in rice: recent progress and future perspectives,” in Molecular breeding for rice abiotic stress tolerance and nutritional quality (Hoboken, New Jersey, USA: John Wiley & Sons, Ltd). doi: 10.1002/9781119633174.ch5

Crossref Full Text | Google Scholar

Cai, Z., He, F., Feng, X., Liang, T., Wang, H., Ding, S., et al. (2020). Transcriptomic analysis reveals important roles of lignin and flavonoid biosynthetic pathways in rice thermotolerance during reproductive stage. Front. Genet. 11. doi: 10.3389/fgene.2020.562937

PubMed Abstract | Crossref Full Text | Google Scholar

Cai, Z., He, F., Feng, X., Liang, T., Wang, H., Ding, S., et al. (2023). Time-series transcriptomic analysis of contrasting rice materials under heat stress reveals a faster response in the tolerant cultivar. Int. J. Mol. Sci. 24, 9408. doi: 10.3390/ijms24119408

PubMed Abstract | Crossref Full Text | Google Scholar

Cao, Y. Y., Chen, Y. H., Chen, M. X., Wang, Z. Q., Wu, C. F., Bian, X. C., et al. (2016). Growth characteristics and endosperm structure of superior and inferior spikelets ofindicarice under high-temperature stress. Biol. Plantarum 60, 3. doi: 10.1007/s10535-016-0606-6

Crossref Full Text | Google Scholar

Cao, Z., Li, Y., Tang, H., Zeng, B., Tang, X., Long, Q., et al. (2020). Fine Mapping of the qHTB1-1QTL, Which Confers Heat Tolerance at the Booting Stage, Using an Oryza Rufipogon Griff. Introgression Line. Theor. Appl. Genet. 133, 1161–1175. doi: 10.1007/s00122-020-03539-7

PubMed Abstract | Crossref Full Text | Google Scholar

Cao, Z., Tang, H., Cai, Y., Zeng, B., Zhao, J., Tang, X., et al. (2022). Natural variation of HTH5 from wild rice, oryza rufipogon griff., is involved in conferring high-temperature tolerance at the heading stage. Plant Biotechnol. J. 20, 1591–1605. doi: 10.1111/pbi.13835

PubMed Abstract | Crossref Full Text | Google Scholar

Ceccarelli, S., Grando, C., Maatougui, M., Michael, M., Slash, M., Haghparast, R., et al. (2010). Plant breeding and climate changes. J. Agric. Sci. 148, 627–637. doi: 10.1017/S0021859610000651

Crossref Full Text | Google Scholar

Chakraborty, A. and Bhattacharjee, S. (2015). Differential competence of redox-regulatory mechanism under extremes of temperature determines growth performances and cross tolerance in two indica rice cultivars. J. Plant Physiol. 176, 65–77. doi: 10.1016/j.jplph.2014.10.016

PubMed Abstract | Crossref Full Text | Google Scholar

Chang, Y., Nguyen, B. H., Xie, Y., Xiao, B., Tang, N., Zhu, W., et al. (2017). Co-overexpression of the constitutively active form of osbZIP46 and ABA-activated protein kinase SAPK6 improves drought and temperature stress resistance in rice. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01102

PubMed Abstract | Crossref Full Text | Google Scholar

Chaudhary, C., Sharma, N., and Khurana, P. (2021). Decoding the wheat awn transcriptome and overexpressing taRca1β in rice for heat stress tolerance. Plant Mol. Biol. 105, 133–146. doi: 10.1007/s11103-020-01073-0

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, C., Begcy, K., Liu, K., Folsom, J. J., Wang, Z., Zhang, C., et al. (2016). Heat stress yields a unique MADS box transcription factor in determining seed size and thermal sensitivity1[OPEN. Plant Physiol. 171, 606–622. doi: 10.1104/pp.15.01992

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, J. H., Chen, S. T., He, N. Y., Wang, Q. L., Zhao, Y., Gao, W., et al. (2020). Nuclear-encoded synthesis of the D1 subunit of photosystem II increases photosynthetic efficiency and crop yield. Nat. Plants 6, 570–580. doi: 10.1038/s41477-020-0629-z

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, F., Dong, G., Wang, F., Shi, Y., Zhu, J., Zhang, Y., et al. (2021). A B-ketoacyl carrier protein reductase confers heat tolerance via the regulation of fatty acid biosynthesis and stress signaling in rice. New Phytol. 232, 655–672. doi: 10.1111/nph.17619

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, M., Fu, Y., Mou, Q., An, J., Zhu, X., Ahmed, T., et al. (2021). Spermidine induces expression of stress associated proteins (SAPs) genes and protects rice seed from heat stress-induced damage during grain-filling. Antioxidants 10, 10. doi: 10.3390/antiox10101544

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, K., Guo, T., Li, X. M., Zhang, Y. M., Yang, Y. B., Ye, W. W., et al. (2019). Translational regulation of plant response to high temperature by a dual-function tRNAHis guanylyltransferase in rice. Mol. Plant 12, 1123–1142. doi: 10.1016/j.molp.2019.04.012

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, J., Tang, L., Shi, P., Yang, B., Sun, T., Cao, W., et al. (2017). Effects of short-term high temperature on grain quality and starch granules of rice (Oryza sativa L.) at post-anthesis stage. Protoplasma 254, 935–943. doi: 10.1007/s00709-016-1002-y

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, L., Wang, Q., Tang, M., Zhang, X., Pan, Y., Yang, X., et al. (2021). QTL mapping and identification of candidate genes for heat tolerance at the flowering stage in rice. Front. Genet. 11, 655–672. doi: 10.3389/fgene.2020.621871

PubMed Abstract | Crossref Full Text | Google Scholar

Chen, Z., Zhou, W., Guo, X., Ling, S., Wang, L., Xin, W., et al. (2024). Heat stress responsive aux/IAA protein, osIAA29 regulates grain filling through osARF17 mediated auxin signaling pathway. Rice 17, 16. doi: 10.1186/s12284-024-00694-z

PubMed Abstract | Crossref Full Text | Google Scholar

Cheng, L. R., Wang, J. M., Uzokwe, V., Meng, L. J., Wang, Y., Sun, Y., et al. (2012). Genetic Analysis of Cold Tolerance at Seedling Stage and Heat Tolerance at Anthesis in Rice (Oryza Sativa L.). J. Integr. Agric. 3, 359–367. doi: 10.1016/S2095-3119(12)60020-3

Crossref Full Text | Google Scholar

Cheng, Q., Zhou, Y., Liu, Z., Zhang, L., Song, G., Guo, Z., et al. (2015). An alternatively spliced heat shock transcription factor, osHSFA2dI, functions in the heat stress-induced unfolded protein response in rice. Plant Biol. (Stuttgart Germany) 17, 419–429. doi: 10.1111/plb.12267

PubMed Abstract | Crossref Full Text | Google Scholar

Choi, J., Choudhury, A. R., Walitang, D. I., Lee, Y., and Sa, T. (2022). “ACC deaminase-producing brevibacterium linens RS16 enhances heat-stress tolerance of rice (Oryza sativa L.). Physiologia Plantarum 174, e135845. doi: 10.1111/ppl.13584

PubMed Abstract | Crossref Full Text | Google Scholar

Cline, W. R. (2007). Global warming and agriculture (New York City: Columbia University Press).

Google Scholar

Coast, O., Murdoch, A. J., Ellis, R. H., Hay, F. R., and Jagadish, K. S. V. (2016). “Resilience of rice (Oryza spp.) pollen germination and tube growth to temperature stress. Plant Cell Environ. 39, 26–375. doi: 10.1111/pce.12475

PubMed Abstract | Crossref Full Text | Google Scholar

Correia, S. d L., Laosuntisuk, K., Desai, J., Ferreira da Silva, P. R., Delatorre, C. A., and Doherty, C. J. (2025). Physiological and molecular responses to high-temperature stress at anthesis in Brazilian flooded rice. AoB Plants 17, plaf043. doi: 10.1093/aobpla/plaf043

PubMed Abstract | Crossref Full Text | Google Scholar

Costa, M. V. J. D., Ramegowda, Y., Ramegowda, V., Karaba, N. N., Sreeman, S. M., and Udayakumar, M. (2021). “Combined drought and heat stress in rice: responses, phenotyping and strategies to improve tolerance. Rice Sci. 28, 233–425. doi: 10.1016/j.rsci.2021.04.003

Crossref Full Text | Google Scholar

Crisp, P., Ganguly, D., Eichten, S., Borevitz, J., and Pogson, B. (2016). Reconsidering plant memory: intersections between stress recovery, RNA turnover, and epigenetics. Sci. Adv. 2, e1501340–e1501340. doi: 10.1126/sciadv.1501340

PubMed Abstract | Crossref Full Text | Google Scholar

Cui, Y., Lu, S., Li, Z., Cheng, J., Hu, P., Zhu, T., et al. (2020). CYCLIC NUCLEOTIDE-GATED ION CHANNELs 14 and 16 promote tolerance to heat and chilling in rice. Plant Physiol. 183, 1794–1808. doi: 10.1104/pp.20.00591

PubMed Abstract | Crossref Full Text | Google Scholar

Das, S., Krishnan, P., Nayak, M., and Ramakrishnan, B. (2014). High temperature stress effects on pollens of rice (Oryza sativa L.) genotypes. Environ. Exp. Bot. 101, 36–46. doi: 10.1016/j.envexpbot.2014.01.004

Crossref Full Text | Google Scholar

Das, P., Lakra, N., Nutan, K. K., Singla-Pareek, S. L., and Pareek, A. (2019). “A unique bZIP transcription factor imparting multiple stress tolerance in rice. Rice 12, 585. doi: 10.1186/s12284-019-0316-8

PubMed Abstract | Crossref Full Text | Google Scholar

Das, A., Pal, M., Taria, S., Ellur, R. K., Kumar, S., Sathee, L., et al. (2024). Multivariate Analysis and Genome Wide Association Mapping for Chlorophyll Fluorescence and Grain Number per Panicle under High Temperature Stress in Rice. Plant Physiol. Rep. 36–46. doi: 10.1007/s40502-024-00808-1

Crossref Full Text | Google Scholar

Deng, Y., Zhu, Z., Chen, J., Kuang, L., Yan, T., Wu, D., et al. (2025). Comparative transcriptomics of indica and japonica rice roots under heat stress reveals the crucial role of osMAPK3 in heat response. Plant Physiol. Biochem. 221, 109668. doi: 10.1016/j.plaphy.2025.109668

PubMed Abstract | Crossref Full Text | Google Scholar

De Storme, N. and Geelen, D. (2014). The impact of environmental stress on male reproductive development in plants: biological processes and molecular mechanisms. Plant Cell Environ. 37, 1–185. doi: 10.1111/pce.12142

PubMed Abstract | Crossref Full Text | Google Scholar

Dhanda, S. S. and Munjal, R. (2012). Heat tolerance in relation to acquired thermotolerance for membrane lipids in bread wheat. Field Crops Res. 135, 30–37. doi: 10.1016/j.fcr.2012.06.009

Crossref Full Text | Google Scholar

Dingkuhn, M., Pasco, R., Pasuquin, J. M., Damo, J., Soulié, J. C., Raboin, L. M., et al. (2017). Crop-model assisted phenomics and genome-wide association study for climate adaptation of indica rice. 2. Thermal stress and spikelet sterility. J. Exp. Bot. 68, 4389–4406. doi: 10.1093/jxb/erx250

PubMed Abstract | Crossref Full Text | Google Scholar

Do, J.-M., Kim, H.-J., Shin, S.-Y., Park, S.-I., Kim, J.-J., and Yoon, H.-S. (2023). “OsHSP 17.9, a small heat shock protein, confers improved productivity and tolerance to high temperature and salinity in a natural paddy field in transgenic rice plants. Agriculture 13, 55. doi: 10.3390/agriculture13050931

Crossref Full Text | Google Scholar

Dong, N. Q., Sun, Y., Guo, T., Shi, C. L., Zhang, Y. M., Kan, Y., et al. (2020). UDP-glucosyltransferase regulates grain size and abiotic stress tolerance associated with metabolic flux redirection in rice. Nat. Commun. 11, 2629. doi: 10.1038/s41467-020-16403-5

PubMed Abstract | Crossref Full Text | Google Scholar

Dong, G. R., Zhao, S. M., Ding, Y., Ma, Y. Q., Ma, X. M., Liu, C. L., et al. (2025). Rice glycosyltransferase osDUGT1 is involved in heat stress tolerance by glycosylating flavonoids and regulating flavonoid metabolism. Front. Plant Sci. 15. doi: 10.3389/fpls.2024.1516990

PubMed Abstract | Crossref Full Text | Google Scholar

Dou, Z., Zhou, Y., Zhang, Y., Guo, W., Xu, Q., and Gao, H. (2024). “Influence of Nitrogen Applications during Grain-Filling Stage on Rice (Oryza Sativa L.) Yield and Grain Quality under High Temperature. Agronomy 14, 15. doi: 10.3390/agronomy14010216

Crossref Full Text | Google Scholar

Driedonks, N., Rieu, I., and Vriezen, W. H. (2016). “Breeding for plant heat tolerance at vegetative and reproductive stages. Plant Reprod. 29, 67–795. doi: 10.1007/s00497-016-0275-9

PubMed Abstract | Crossref Full Text | Google Scholar

El-Esawi, M. A. and Alayafi, A. A. (2019). “Overexpression of rice rab7 gene improves drought and heat tolerance and increases grain yield in rice (Oryza sativa L.). Genes 10, 565. doi: 10.3390/genes10010056

PubMed Abstract | Crossref Full Text | Google Scholar

El-kereamy, A., Bi, Y.-M., Ranathunge, K., Beatty, P. H., Good, A. G., and Rothstein, S. J. (2012). “The rice R2R3-MYB transcription factor osMYB55 is involved in the tolerance to high temperature and modulates amino acid metabolism. PloS One 7, e520305. doi: 10.1371/journal.pone.0052030

PubMed Abstract | Crossref Full Text | Google Scholar

Endo, M., Tsuchiya, T., Hamada, K., Kawamura, S., Yano, K., Ohshima, M., et al. (2009). High temperatures cause male sterility in rice plants with transcriptional alterations during pollen development. Plant Cell Physiol. 50, 1911–1922. doi: 10.1093/pcp/pcp135

PubMed Abstract | Crossref Full Text | Google Scholar

Essemine, J., Xiao, Y., Qu, M., Mi, H., and Zhu, X.-G. (2017). Cyclic Electron Flow May Provide Some Protection against PSII Photoinhibition in Rice (Oryza Sativa L.) Leaves under Heat Stress. J. Plant Physiol. 211, 138–146. doi: 10.1016/j.jplph.2017.01.007

PubMed Abstract | Crossref Full Text | Google Scholar

Fahad, S., Bajwa, A. A., Nazir, U., Anjum, S. A., Farooq, A., Zohaib, A., et al. (2017). Crop production under drought and heat stress: plant responses and management options. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01147

PubMed Abstract | Crossref Full Text | Google Scholar

Fahad, S., Hussain, S., Saud, S., Hassan, S., Ihsan, Z., Shah, A. N., et al. (2016). Exogenously applied plant growth regulators enhance the morpho-physiological growth and yield of rice under high temperature. Front. Plant Sci. 7. doi: 10.3389/fpls.2016.01250

PubMed Abstract | Crossref Full Text | Google Scholar

Fang, Y., Liao, K., Du, H., Xu, Y., Song, H., Li, X., et al. (2015). A stress-responsive NAC transcription factor SNAC3 confers heat and drought tolerance through modulation of reactive oxygen species in rice. J. Exp. Bot. 66, 6803–6817. doi: 10.1093/jxb/erv386

PubMed Abstract | Crossref Full Text | Google Scholar

FAOSTAT. Available online at: https://www.fao.org/faostat/en/data/QCL (Accessed February 10, 2025).

Google Scholar

Feng, L., Wang, K., Li, Y., Tan, Y., Kong, J., Li, H., et al. (2007). Overexpression of SBPase enhances photosynthesis against high temperature stress in transgenic rice plants. Plant Cell Rep. 26, 1635–1646. doi: 10.1007/s00299-006-0299-y

PubMed Abstract | Crossref Full Text | Google Scholar

Feng, B., Xu, Y., Fu, W., Li, H., Li, G., Li, J., et al. (2023). RGA1 negatively regulates thermo-tolerance by affecting carbohydrate metabolism and the energy supply in rice. Rice 16, 32. doi: 10.1186/s12284-023-00649-w

PubMed Abstract | Crossref Full Text | Google Scholar

Feng, B., Zhang, C., Chen, T., Zhang, X., Tao, L., and Fu, G. (2018). “Salicylic acid reverses pollen abortion of rice caused by heat stress. BMC Plant Biol. 18, 2455. doi: 10.1186/s12870-018-1472-5

PubMed Abstract | Crossref Full Text | Google Scholar

Fengfeng, F., Meng, C., Xiong, L., Manman, L., Huanran, Y., Mingxing, C., et al. (2023). Novel QTLs from wild rice oryza longistaminata confer strong tolerance to high temperature at seedling stage. Rice Sci. 30, 577–586. doi: 10.1016/j.rsci.2023.07.004

Crossref Full Text | Google Scholar

Finkelstein, R., Reeves, W., Ariizumi, T., and Steber, C. (2008). Molecular aspects of seed dormancy*. Annu. Rev. Plant Biol. 59, 387–415. doi: 10.1146/annurev.arplant.59.032607.092740

PubMed Abstract | Crossref Full Text | Google Scholar

Firon, N., Shaked, R., Peet, M. M., Pharr, D. M., Zamski, E., Rosenfeld, K., et al. (2006). Pollen grains of heat tolerant tomato cultivars retain higher carbohydrate concentration under heat stress conditions. Scientia Hortic. 109, 212–217. doi: 10.1016/j.scienta.2006.03.007

Crossref Full Text | Google Scholar

Fitzgerald, M. A., McCouch, S. R., and Hall, R. D. (2009). “Not just a grain of rice: the quest for quality. Trends Plant Sci. 14, 133–395. doi: 10.1016/j.tplants.2008.12.004

PubMed Abstract | Crossref Full Text | Google Scholar

Folsom, J. J., Begcy, K., Hao, X., Wang, D., and Walia, H. (2014). “Rice fertilization-independent endosperm1 regulates seed size under heat stress by controlling early endosperm development. Plant Physiol. 165, 238–485. doi: 10.1104/pp.113.232413

PubMed Abstract | Crossref Full Text | Google Scholar

Fu, G., Feng, B., Zhang, C., Yang, Y., Yang, X., Chen, T., et al. (2016). Heat stress is more damaging to superior spikelets than inferiors of rice (Oryza sativa L.) due to their different organ temperatures. Front. Plant Sci. 7. doi: 10.3389/fpls.2016.01637

PubMed Abstract | Crossref Full Text | Google Scholar

Gammulla, C., Pascovici, D., Atwell, B., and Haynes, P. (2010). Differential metabolic response of cultured rice (Oryza sativa) cells exposed to high- and low-temperature stress. Proteomics 10, 3001–3019. doi: 10.1002/pmic.201000054

PubMed Abstract | Crossref Full Text | Google Scholar

Gandass, N., Kajal, and Salvi, P. (2022). Intrinsically disordered protein, DNA binding with one finger transcription factor (OsDOF27) implicates thermotolerance in yeast and rice. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.956299

PubMed Abstract | Crossref Full Text | Google Scholar

Gardiner, L.-J., Quinton-Tulloch, M., Olohan, L., Price, J., Hall, N., and Hall, A. (2015). “A genome-wide survey of DNA methylation in hexaploid wheat. Genome Biol. 16, 2735. doi: 10.1186/s13059-015-0838-3

PubMed Abstract | Crossref Full Text | Google Scholar

Gautam, H., Fatma, M., Sehar, Z., Iqbal, N., Albaqami, M., and Khan., N. A. (2022). “Exogenously-sourced ethylene positively modulates photosynthesis, carbohydrate metabolism, and antioxidant defense to enhance heat tolerance in rice. Int. J. Mol. Sci. 23, 35. doi: 10.3390/ijms23031031

PubMed Abstract | Crossref Full Text | Google Scholar

Gill, T., Gill, S. K., Saini, D. K., Chopra, Y., de Koff, J. P., and Sandhu, K. S. (2022). “A comprehensive review of high throughput phenotyping and machine learning for plant stress phenotyping. Phenomics 2, 1565. doi: 10.1007/s43657-022-00048-z

PubMed Abstract | Crossref Full Text | Google Scholar

Gong, Z., Xiong, L., Shi, H., Yang, S., Herrera-Estrella, L. R., Xu, G., et al. (2020). Plant abiotic stress response and nutrient use efficiency. Sci. China Life Sci. 63, 635–674. doi: 10.1007/s11427-020-1683-x

PubMed Abstract | Crossref Full Text | Google Scholar

Gonzalez, C., Maria, C., Cristo, E., Pérez, N., Reyes, Y., Horta, D., et al. (2021). Evaluation of mutant rice genotypes for tolerance to high temperature. Aust. J. Crop Sci. 15, 43–49. doi: 10.3316/informit.176336772423923

Crossref Full Text | Google Scholar

Gourdji, S. M., Sibley, A. M., and Lobell, D. B. (2013). “Global crop exposure to critical high temperatures in the reproductive period: historical trends and future projections. Environ. Res. Lett. 8, 0240415. doi: 10.1088/1748-9326/8/2/024041

Crossref Full Text | Google Scholar

Gu, X., Si, F., Feng, Z., Li, S., Liang, D., Yang, P., et al. (2023). The osSGS3-tasiRNA-osARF3 module orchestrates abiotic-biotic stress response trade-off in rice. Nat. Commun. 14, 4441. doi: 10.1038/s41467-023-40176-2

PubMed Abstract | Crossref Full Text | Google Scholar

Guan, Y., Chen, Y., Huang, Q., He, Y., Li, X., Zhu, Z., et al. (2025). Exploring heat stress responses and heat tolerance in rice in the reproductive stage: A dual omics approach. Plant Growth Regul. 105, 1459–1477. doi: 10.1007/s10725-025-01352-0

Crossref Full Text | Google Scholar

Guan-fu, F. U., Long-xing, T. A. O., Jian, S., Xi, W., Li-yong, C., and Shi-hua, C. (2008). Responses of yield characteristics to high temperature during flowering stage in hybrid rice guodao 6. Rice Sci. 15, 2155.

Google Scholar

Guihur, A., Fauvet, B., Finka, A., Quadroni, M., and Goloubinoff, P. (2021). “Quantitative proteomic analysis to capture the role of heat-accumulated proteins in moss plant acquired thermotolerance. Plant Cell Environ. 44, 2117–2335. doi: 10.1111/pce.13975

PubMed Abstract | Crossref Full Text | Google Scholar

Guo, H., Tao, W., Gao, H. Y., Chen, L., Zhong, X. Y., Tang, M., et al. (2024). Physiological traits, gene expression responses, and proteomics of rice varieties varying in heat stress tolerance at the flowering stage. Front. Plant Sci. 15. doi: 10.3389/fpls.2024.1489331

PubMed Abstract | Crossref Full Text | Google Scholar

Guo, M., Zhang, X., Liu, J., Hou, L., Liu, H., and Zhao, X. (2020). “OsProDH negatively regulates thermotolerance in rice by modulating proline metabolism and reactive oxygen species scavenging. Rice 13, 615. doi: 10.1186/s12284-020-00422-3

PubMed Abstract | Crossref Full Text | Google Scholar

Haider, S., Iqbal, J., Naseer, S., Yaseen, T., Shaukat, M., Bibi, H., et al. (2021). Molecular mechanisms of plant tolerance to heat stress: current landscape and future perspectives. Plant Cell Rep. 40, 2247–2271. doi: 10.1007/s00299-021-02696-3

PubMed Abstract | Crossref Full Text | Google Scholar

Harshada, V.K., Patil Arun, H., Chaudhari, P., and Chandel, G. (2021). Impact of heat stress on expression pattern of nine rice heat shock factor genes and its traits related to tolerance. Indian J. Biotechnol. 20, 65–75.

Google Scholar

Hasanuzzaman, M., Nahar, K., Alam, M., Roychowdhury, R., and Fujita, M. (2013). “Physiological, biochemical, and molecular mechanisms of heat stress tolerance in plants. Int. J. Mol. Sci. 14, 9643–9845. doi: 10.3390/ijms14059643

PubMed Abstract | Crossref Full Text | Google Scholar

Hasegawa, T., Ishimaru, T., Kondo, M., Kuwagata, T., Yoshimoto, M., and Fukuoka, M. (2011). Spikelet sterility of rice observed in the record hot summer of 2007 and the factors associated with its variation. Journal of Agricultural Meteorology 67, 225–232. doi: 10.2480/agrmet.67.4.3

Crossref Full Text | Google Scholar

Hassan, M. U., Chattha, M. U., Khan, I., Chattha, M. B., Barbanti, L., Aamer, M., et al. (2020). Heat stress in cultivated plants: nature, impact, mechanisms, and mitigation strategies - A review. Plant Biosyst. - Int. J. Dealing All Aspects Plant Biol. 155, 1–56. doi: 10.1080/11263504.2020.1727987

Crossref Full Text | Google Scholar

He, Y., Guan, H., Li, B., Zhang, S., Xu, Y., Yao, Y., et al. (2023). Transcriptome analysis reveals the dynamic and rapid transcriptional reprogramming involved in heat stress and identification of heat response genes in rice. Int. J. Mol. Sci. 24, 14802. doi: 10.3390/ijms241914802

PubMed Abstract | Crossref Full Text | Google Scholar

He, Y., Zhang, X., Shi, Y., Xu, X., Li, L., and Wu, J.-L. (2021). “Premature senescence leaf 50 Promotes Heat Stress Tolerance in Rice (Oryza Sativa L.). Rice 14, 535. doi: 10.1186/s12284-021-00493-w

PubMed Abstract | Crossref Full Text | Google Scholar

Higashi, Y. and Saito, K. (2019). Lipidomic studies of membrane glycerolipids in plant leaves under heat stress. Prog. Lipid Res. 75, 100990. doi: 10.1016/j.plipres.2019.100990

PubMed Abstract | Crossref Full Text | Google Scholar

Hirabayashi, H., Sasaki, K., Kambe, T., Gannaban, R. B., Miras, M. A., Mendioro, M. S., et al. (2015). QEMF3, a Novel QTL for the Early-Morning Flowering Trait from Wild Rice, Oryza Officinalis, to Mitigate Heat Stress Damage at Flowering in Rice, O. Sativa. J. Exp. Bot. 66, 1227–1236. doi: 10.1093/jxb/eru474

PubMed Abstract | Crossref Full Text | Google Scholar

Hoang, T. V., Vo, K. T. X., Rahman, M. M., Choi, S.-H., and Jeon, J.-S. (2019). Heat stress transcription factor osSPL7 plays a critical role in reactive oxygen species balance and stress responses in rice. Plant Sci. 289, 110273. doi: 10.1016/j.plantsci.2019.110273

PubMed Abstract | Crossref Full Text | Google Scholar

Hosseiniyan Khatibi, S. M., Dimaano, N. G., Veliz, E., Sundaresan, V., and Ali, J. (2024). “Exploring and exploiting the rice phytobiome to tackle climate change challenges. Plant Commun. 5 (12), 101078. doi: 10.1016/j.xplc.2024.101078

PubMed Abstract | Crossref Full Text | Google Scholar

Hu, S., Ding, Y., and Zhu, C. (2020). Sensitivity and responses of chloroplasts to heat stress in plants. Front. Plant Sci. 11. doi: 10.3389/fpls.2020.00375

PubMed Abstract | Crossref Full Text | Google Scholar

Hu, C., Jiang, J., Li, Y., Song, S., Zou, Y., Jing, C., et al. (2022). QTL mapping and identification of candidate genes using a genome-wide association study for heat tolerance at anthesis in rice (Oryza sativa L.). Front. Genet. 13. doi: 10.3389/fgene.2022.983525

PubMed Abstract | Crossref Full Text | Google Scholar

Hu, Q., Wang, W., Lu, Q., Huang, J., Peng, S., and Cui, K. (2021). “Abnormal Anther Development Leads to Lower Spikelet Fertility in Rice (Oryza Sativa L.) under High Temperature during the Panicle Initiation Stage. BMC Plant Biol. 21, 4285. doi: 10.1186/s12870-021-03209-w

PubMed Abstract | Crossref Full Text | Google Scholar

Huang, S., Aken, O. V., Schwarzländer, M., Belt, K., and Millar, A.H. (2016). “The roles of mitochondrial reactive oxygen species in cellular signaling and stress response in plants. Plant Physiol. 171, 1551–1595. doi: 10.1104/pp.16.00166

PubMed Abstract | Crossref Full Text | Google Scholar

Huang, D., Zhang, Z., Fan, Y., Tang, S., Zhuang, J., and Zhu, Y. (2023). “Detection of QTL for high-temperature tolerance in rice using a high-density bin map. Agronomy 13, 65. doi: 10.3390/agronomy13061582

Crossref Full Text | Google Scholar

Huang, M., Zhang, H., Zhao, C., Chen, G., and Zou, Y. (2019). “Amino acid content in rice grains is affected by high temperature during the early grain-filling period. Sci. Rep. 9, 27005. doi: 10.1038/s41598-019-38883-2

PubMed Abstract | Crossref Full Text | Google Scholar

Hwang, O. J. and Back, K. (2019). “Melatonin deficiency confers tolerance to multiple abiotic stresses in rice via decreased brassinosteroid levels. Int. J. Mol. Sci. 20, 51735. doi: 10.3390/ijms20205173

PubMed Abstract | Crossref Full Text | Google Scholar

Hwang, S. K., Singh, S., Maharana, J., Kalita, S., Tuncel, A., Rath, T., et al. (2019). Mechanism underlying heat stability of the rice endosperm cytosolic ADP-glucose pyrophosphorylase. Front. Plant Sci. 10. doi: 10.3389/fpls.2019.00070

PubMed Abstract | Crossref Full Text | Google Scholar

Impa, S. M., Raju, B., Hein, N. T., Sandhu, J., Prasad, P. V. V., Walia, H., et al. (2021). High night temperature effects on wheat and rice: current status and way forward. Plant Cell Environ. 44, 2049–2065. doi: 10.1111/pce.14028

PubMed Abstract | Crossref Full Text | Google Scholar

IPCC AR5 climate change 2014: mitigation of climate change — IPCC. Available online at: https://www.ipcc.ch/report/ar5/wg3/ (Accessed April 5, 2025).

Google Scholar

Iqbal, Z., Iqbal, M. S., Khan, M.I. R., and Ansari, M. I. (2021). “Toward integrated multi-omics intervention: rice trait improvement and stress management. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.741419

PubMed Abstract | Crossref Full Text | Google Scholar

Ishimaru, T., Hirabayashi, H., Ida, M., Takai, T., San-Oh, Y. A., Yoshinaga, S., et al. (2010). A genetic resource for early-morning flowering trait of wild rice oryza officinalis to mitigate high temperature-induced spikelet sterility at anthesis. Ann. Bot. 106, 515–520. doi: 10.1093/aob/mcq124

PubMed Abstract | Crossref Full Text | Google Scholar

Jagadish, S.V.K. (2020). “Heat stress during flowering in cereals - effects and adaptation strategies. New Phytol. 226, 1567–1725. doi: 10.1111/nph.16429

PubMed Abstract | Crossref Full Text | Google Scholar

Jagadish, S. V. K., Craufurd, P. Q., and Wheeler, T. R. (2007). High temperature stress and spikelet fertility in rice (Oryza sativa L.). J. Exp. Bot. 58, 1627–1635. doi: 10.1093/jxb/erm003

PubMed Abstract | Crossref Full Text | Google Scholar

Jagadish, S. V. K., Murty, M. V. R., and Quick, W. P. (2015). Rice responses to rising temperatures–challenges, perspectives and future directions. Plant Cell Environ. 38, 1686–1698. doi: 10.1111/pce.12430

PubMed Abstract | Crossref Full Text | Google Scholar

Jagadish, S. V. K., Muthurajan, R., Oane, R., Wheeler, T. R., Heuer, S., Bennett, J., et al. (2010). Physiological and proteomic approaches to address heat tolerance during anthesis in rice (Oryza sativa L.). J. Exp. Bot. 61, 143–156. doi: 10.1093/jxb/erp289

PubMed Abstract | Crossref Full Text | Google Scholar

Jan, R., Kim, N., Lee, S. H., Khan, M. A., Asaf, S., Lubna., et al. (2021). Enhanced flavonoid accumulation reduces combined salt and heat stress through regulation of transcriptional and hormonal mechanisms. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.796956

PubMed Abstract | Crossref Full Text | Google Scholar

Janni, M., Gullì, M., Maestri, E., Marmiroli, M., Valliyodan, B., Nguyen, H. T., et al. (2020). Molecular and genetic bases of heat stress responses in crop plants and breeding for increased resilience and productivity. J. Exp. Bot. 71, 3780–3802. doi: 10.1093/jxb/eraa034

PubMed Abstract | Crossref Full Text | Google Scholar

Jeong, E. J., Song, J. Y., Yu, D. A., Kim, M. S., Jung, Y. J., Kang, K. Q., et al. (2017). Overexpression of an oligopeptide transporter gene enhances heat tolerance in transgenic rice. J. Plant Biotechnol. 44, 296–302. doi: 10.5010/JPB.2017.44.3.296

Crossref Full Text | Google Scholar

Jhan, L. H., Yang, C. Y., Huang, C. M., Lai, M. C., Huang, Y. H., Baiya, S., et al. (2023). Integrative pathway and network analysis provide insights on flooding-tolerance genes in soybean. Sci. Rep. 13, 1980. doi: 10.1038/s41598-023-28593-1

PubMed Abstract | Crossref Full Text | Google Scholar

Jiang, Z., Tu, H., Bai, B., Yang, C., Zhao, B., Guo, Z., et al. (2021). Combining UAV-RGB high-throughput field phenotyping and genome-wide association study to reveal genetic variation of rice germplasms in dynamic response to drought stress. New Phytol. 232, 440–455. doi: 10.1111/nph.17580

PubMed Abstract | Crossref Full Text | Google Scholar

Jones, M. W., Peters, G. P., Gasser, T., Andrew, R. M., Schwingshackl, C., Gütschow, J., et al. (2024). National contributions to climate change due to historical emissions of carbon dioxide, methane and nitrous oxide. Zenodo, 155. doi: 10.5281/zenodo.14054503

PubMed Abstract | Crossref Full Text | Google Scholar

Ju, Y.-H., Choi, J.-Y., Yun, S.-H., Mittra, P. K., Woo, S.-H., and Sakagami, J.-I. (2025). “Stage-dependent heat priming mitigates reproductive heat stress via proteomic regulation in oryza sativa L. Plant Sci. 359, 112639. doi: 10.1016/j.plantsci.2025.112639

PubMed Abstract | Crossref Full Text | Google Scholar

Kan, Y. and Lin, H.-X. (2021). Molecular regulation and genetic control of rice thermal response. Crop Journal Rice as Model. crop: genetics Genomics Breed. 9, 497–505. doi: 10.1016/j.cj.2021.02.008

Crossref Full Text | Google Scholar

Kan, Y., Mu, X.-R., Gao, J., Lin, H.-X., and Lin, Y. (2023). “The molecular basis of heat stress responses in plants. Mol. Plant 16, 1612–1345. doi: 10.1016/j.molp.2023.09.013

PubMed Abstract | Crossref Full Text | Google Scholar

Kan, Y., Mu, X. R., Zhang, H., Gao, J., Shan, J. X., Ye, W. W., et al. (2022). TT2 controls rice thermotolerance through SCT1-dependent alteration of wax biosynthesis. Nat. Plants 8, 53–67. doi: 10.1038/s41477-021-01039-0

PubMed Abstract | Crossref Full Text | Google Scholar

Kang, Z., Qin, T., and Zhao, Z. (2019). “Overexpression of the zinc finger protein gene osZFP350 improves root development by increasing resistance to abiotic stress in rice. Acta Biochim. Polonica 66, 25. doi: 10.18388/abp.2018_2765

PubMed Abstract | Crossref Full Text | Google Scholar

Karim, M. R., Ishikawa, M., Ikeda, M., and Islam, T. (2012). “Climate change model predicts 33 % Rice yield decrease in 2100 in Bangladesh. Agron. Sustain. Dev. 32, 821–305. doi: 10.1007/s13593-012-0096-7

Crossref Full Text | Google Scholar

Katiyar-Agarwal, S., Agarwal, M., and Grover, A. (2003). “Heat-tolerant basmati rice engineered by over-expression of hsp101. Plant Mol. Biol. 51, 677–865. doi: 10.1023/a:1022561926676

PubMed Abstract | Crossref Full Text | Google Scholar

Khan, S., Anwar, S., Ashraf, M. Y., Khaliq, B., Sun, M., Hussain, S., et al. (2019). Mechanisms and adaptation strategies to improve heat tolerance in rice. A review. Plants 8, 508. doi: 10.3390/plants8110508

PubMed Abstract | Crossref Full Text | Google Scholar

Khan, M., Iqbal, R., Kumari, S., Nazir, F., Khanna, R. R., Gupta, R., et al. (2023). “Defensive role of plant hormones in advancing abiotic stress-resistant rice plants. Rice Sci. 30, 15–355. doi: 10.1016/j.rsci.2022.08.002

Crossref Full Text | Google Scholar

Khatibi, S. M. H. and Ali, J. (2024). Harnessing the power of machine learning for crop improvement and sustainable production. Front. Plant Sci. 15. doi: 10.3389/fpls.2024.1417912

PubMed Abstract | Crossref Full Text | Google Scholar

Kilasi, N. L., Singh, J., Vallejos, C. E., Ye, C., Jagadish, S. V. K., Kusolwa, P., et al. (2018). Heat stress tolerance in rice (Oryza sativa L.): identification of quantitative trait loci and candidate genes for seedling growth under heat stress. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01578

PubMed Abstract | Crossref Full Text | Google Scholar

Kim, J. H., Lim, S. D., and Jang, C. S. (2019). “Oryza sativa heat-induced RING finger protein 1 (OsHIRP1) positively regulates plant response to heat stress. Plant Mol. Biol. 99, 545–595. doi: 10.1007/s11103-019-00835-9

PubMed Abstract | Crossref Full Text | Google Scholar

Kim, J. H., Lim, S. D., and Jang, C. S. (2020). “Oryza sativa drought-, heat-, and salt-induced RING finger protein 1 (OsDHSRP1) negatively regulates abiotic stress-responsive gene expression. Plant Mol. Biol. 103, 235–525. doi: 10.1007/s11103-020-00989-x

PubMed Abstract | Crossref Full Text | Google Scholar

Kinoshita, T. and Seki, M. (2014). “Epigenetic memory for stress response and adaptation in plants. Plant Cell Physiol. 55, 1859–1635. doi: 10.1093/pcp/pcu125

PubMed Abstract | Crossref Full Text | Google Scholar

Kobayashi, K., Matsui, T., Murata, Y., and Yamamoto, M. (2011). “Percentage of dehisced thecae and length of dehiscence control pollination stability of rice cultivarsat high temperatures. Plant Production Sci. 14, 89–955. doi: 10.1626/pps.14.89

Crossref Full Text | Google Scholar

Korres, N. E., Norsworthy, J. K., Burgos, N. R., and Oosterhuis, D. M. (2017). Temperature and drought impacts on rice production: an agronomic perspective regarding short- and long-term adaptation measures. Water Resour. Rural Dev. 9, 12–27. doi: 10.1016/j.wrr.2016.10.001

Crossref Full Text | Google Scholar

Krishnan, P., Ramakrishnan, B., Reddy, K., and Reddy, V. (2011). High-temperature effects on rice growth, yield, and grain quality. Adv. Agron. - ADVAN Agron. 111, 87–206. doi: 10.1016/B978-0-12-387689-8.00004-7

Crossref Full Text | Google Scholar

Kuang, J., Liu, J., Mei, J., Wang, C., Hu, H., Zhang, Y., et al. (2017). A class II small heat shock protein osHsp18.0 plays positive roles in both biotic and abiotic defense responses in rice. Sci. Rep. 7, 11333. doi: 10.1038/s41598-017-11882-x

PubMed Abstract | Crossref Full Text | Google Scholar

Kumar, A., Dixit, S., Ram, T., Yadaw, R. B., Mishra, K. K., and Mandal, N. P. (2014). “Breeding high-yielding drought-tolerant rice: genetic variations and conventional and molecular approaches. J. Exp. Bot. 65, 6265–6785. doi: 10.1093/jxb/eru363

PubMed Abstract | Crossref Full Text | Google Scholar

Kumar, A., Thomas, J., Gill, N., Dwiningsih, Y., Ruiz, C., Famoso, A., et al. (2023). Molecular mapping and characterization of QTLs for grain quality traits in a RIL population of US rice under high nighttime temperature stress. Sci. Rep. 13, 4880. doi: 10.1038/s41598-023-31399-w

PubMed Abstract | Crossref Full Text | Google Scholar

Kundu, S., Saini, D., Meena, R., Bahuguna, R., and Jagadish, K. (2024). “High-throughput phenotyping and AI technologies for deciphering crop resilience to heat stress. Plant Physiol. Rep. 699–715. doi: 10.1007/s40502-024-00821-4

Crossref Full Text | Google Scholar

Lal, M. K., Tiwari, R. K., Gahlaut, V., Mangal, V., Kumar, A., Singh, M. P., et al. (2022). Physiological and molecular insights on wheat responses to heat stress. Plant Cell Rep. 41, 501–518. doi: 10.1007/s00299-021-02784-4

PubMed Abstract | Crossref Full Text | Google Scholar

Lamers, J., van der Meer, T., and Testerink, C. (2020). “How plants sense and respond to stressful environments. Plant Physiol. 182, 1624–1355. doi: 10.1104/pp.19.01464

PubMed Abstract | Crossref Full Text | Google Scholar

Lang, N. T., Thi Thu Ha, P., Tru, P. C., Toan, T. B., Buu, B. C., and Cho, Y.-C. (2015). “Breeding for heat tolerance rice based on marker-assisted backcrosing in Vietnam. Plant Breed. Biotechnol. 3, 274–815. doi: 10.9787/PBB.2015.3.3.274

Crossref Full Text | Google Scholar

Larkindale, J., Hall, J. D., Knight, M. R., and Vierling, E. (2005). “Heat stress phenotypes of arabidopsis mutants implicate multiple signaling pathways in the acquisition of thermotolerance. Plant Physiol. 138, 882–975. doi: 10.1104/pp.105.062257

PubMed Abstract | Crossref Full Text | Google Scholar

Lawas, L. M. F., Li, X., Erban, A., Kopka, J., Jagadish, S. V. K., Zuther, E., et al. (2019). Metabolic responses of rice cultivars with different tolerance to combined drought and heat stress under field conditions. GigaScience 8, giz050. doi: 10.1093/gigascience/giz050

PubMed Abstract | Crossref Full Text | Google Scholar

Lawas, L. M. F., Zuther, E., Jagadish, S. V. K., and Hincha, D. K. (2018). Molecular mechanisms of combined heat and drought stress resilience in cereals. Curr. Opin. Plant Biology AGRI 2017 45, 212–217. doi: 10.1016/j.pbi.2018.04.002

PubMed Abstract | Crossref Full Text | Google Scholar

Lee, H., Calvin, K., Dasgupta, D., Krinner, G., Mukherji, A., Thorne, P., et al. (2023). IPCC 2023: Climate change 2023: synthesis report. Contribution of Working Groups I, II and III to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change.

Google Scholar

Lee, S. Y. (2002). Germination - varietal differences in seed germination and seedling vigor of korean rice varietics following dry heat treatment. Seed Sci. Technol. 30, 311–322.

Google Scholar

Lee, S.-Y., Kim, Y.-H., and Lee, G.-S. (2017). “Mapping QTLs associated to germination stability following dry-heat treatment in rice seed. 3 Biotech. 7, 2205. doi: 10.1007/s13205-017-0807-5

PubMed Abstract | Crossref Full Text | Google Scholar

Li, B., Cai, H., Liu, K., An, B., Wang, R., Yang, F., et al. (2023). DNA methylation alterations and their association with high temperature tolerance in rice anthesis. J. Plant Growth Regul. 42, 780–794. doi: 10.1007/s00344-022-10586-5

Crossref Full Text | Google Scholar

Li, X. M., Chao, D. Y., Wu, Y., Huang, X., Chen, K., Cui, L. G., et al. (2015). Natural alleles of a proteasome A2 subunit gene contribute to thermotolerance and adaptation of african rice. Nat. Genet. 47, 827–833. doi: 10.1038/ng.3305

PubMed Abstract | Crossref Full Text | Google Scholar

Li, P., Jiang, J., Zhang, G., Miao, S., Lu, J., Qian, Y., et al. (2023). Integrating GWAS and transcriptomics to identify candidate genes conferring heat tolerance in rice. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.1102938

PubMed Abstract | Crossref Full Text | Google Scholar

Li, X., Lawas, L. M. F., Malo, R., Glaubitz, U., Erban, A., Mauleon, R., et al. (2015). Metabolic and transcriptomic signatures of rice floral organs reveal sugar starvation as a factor in reproductive failure under heat and drought stress. Plant Cell Environ. 38, 2171–2192. doi: 10.1111/pce.12545

PubMed Abstract | Crossref Full Text | Google Scholar

Li, Y., Li, J., Chen, Z., Wei, Y., Qi, Y., and Wu, C. (2020). “OsmiR167a-targeted auxin response factors modulate tiller angle via fine-tuning auxin distribution in rice. Plant Biotechnol. J. 18, 2015–2265. doi: 10.1111/pbi.13360

PubMed Abstract | Crossref Full Text | Google Scholar

Li, M. M., Li, X., Yu, L. Q., Wu, J. W., Li, H., Liu, J., et al. (2018). Identification of QTLs associated with heat tolerance at the heading and flowering stage in rice (Oryza sativa L.). Euphytica 214, 70. doi: 10.1007/s10681-018-2136-0

Crossref Full Text | Google Scholar

Li, F., Liu, Y., Zhang, X., Liu, L., Yan, Y., Ji, X., et al. (2022). Transcriptome and metabolome analyses reveals the pathway and metabolites of grain quality under phytochrome B in rice (Oryza sativa L.). Rice 15, 52. doi: 10.1186/s12284-022-00600-5

PubMed Abstract | Crossref Full Text | Google Scholar

Li, Z., Wei, X., Tong, X., Zhao, J., Liu, X., Wang, H., et al. (2022). The osNAC23-tre6P-snRK1a feed-forward loop regulates sugar homeostasis and grain yield in rice. Mol. Plant 15, 706–722. doi: 10.1016/j.molp.2022.01.016

PubMed Abstract | Crossref Full Text | Google Scholar

Li, J., Yang, J., Zhu, B., and Xie, G. (2019). Overexpressing osFBN1 enhances plastoglobule formation, reduces grain-filling percent and jasmonate levels under heat stress in rice. Plant Sci. 285, 230–238. doi: 10.1016/j.plantsci.2019.05.007

PubMed Abstract | Crossref Full Text | Google Scholar

Liang, Z., Zhang, Q., Ji, C., Hu, G., Zhang, P., Wang, Y., et al. (2021). Reorganization of the 3D chromatin architecture of rice genomes during heat stress. BMC Biol. 19, 53. doi: 10.1186/s12915-021-00996-4

PubMed Abstract | Crossref Full Text | Google Scholar

Liao, J.-L., Zhang, H.-Y., Liu, J.-B., Zhong, P.-A., and Huang, Y.-J. (2012). Identification of candidate genes related to rice grain weight under high-temperature stress. Plant Science: Int. J. Exp. Plant Biol. 196, 32–43. doi: 10.1016/j.plantsci.2012.07.013

PubMed Abstract | Crossref Full Text | Google Scholar

Lim, S. D., Cho, H. Y., Park, Y. C., Ham, D. J., Lee, J. K., and Jang, C. S. (2013). “The rice RING finger E3 ligase, osHCI1, drives nuclear export of multiple substrate proteins and its heterogeneous overexpression enhances acquired thermotolerance. J. Exp. Bot. 64, 2899–9145. doi: 10.1093/jxb/ert143

PubMed Abstract | Crossref Full Text | Google Scholar

Lin, M.-y., Chai, K.-h., Ko, S.-s., Kuang, L.-y., Lur, H.-S., and Charng, Y.-y. (2014). “A positive feedback loop between HEAT SHOCK PROTEIN101 and HEAT STRESS-ASSOCIATED 32-KD PROTEIN modulates long-term acquired thermotolerance illustrating diverse heat stress responses in rice varieties. Plant Physiol. 164, 2045–2535. doi: 10.1104/pp.113.229609

PubMed Abstract | Crossref Full Text | Google Scholar

Lin, S., Liu, Z., Sun, S., Xue, F., Li, H., Tursun, A., et al. (2023). Rice HEAT SHOCK PROTEIN60-3B maintains male fertility under high temperature by starch granule biogenesis. Plant Physiol. 192, 2301–2317. doi: 10.1093/plphys/kiad136

PubMed Abstract | Crossref Full Text | Google Scholar

Lin, Y., Zhu, Y., Cui, Y., Qian, H., Yuan, Q., Chen, R., et al. (2023). Identification of natural allelic variation in TTL1 controlling thermotolerance and grain size by a rice super pan-genome. J. Integr. Plant Biol. 65, 2541–2551. doi: 10.1111/jipb.13568

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, J., Hasanuzzaman, M., Wen, H., Zhang, J., Peng, T., Sun, H., et al. (2019). High temperature and drought stress cause abscisic acid and reactive oxygen species accumulation and suppress seed germination growth in rice. Protoplasma 256, 1217–1227. doi: 10.1007/s00709-019-01354-6

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, J.-X. and Howell, S. H. (2016). “Managing the protein folding demands in the endoplasmic reticulum of plants. New Phytol. 211, 418–285. doi: 10.1111/nph.13915

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, X., Ji, P., Liao, J., Duan, X., Luo, Z., Yu, X., et al. (2025). CRISPR/cas knockout of the NADPH oxidase gene osRbohB reduces ROS overaccumulation and enhances heat stress tolerance in rice. Plant Biotechnol. J. 23, 336–351. doi: 10.1111/pbi.14500

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, Y., Liu, X., Wang, X., Gao, K., Wei, W., Ren, H., et al. (2020). Heterologous expression of heat stress-responsive atPLC9 confers heat tolerance in transgenic rice. BMC Plant Biol. 20, 514. doi: 10.1186/s12870-020-02709-5

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, X.-H., Lyu, Y.-S., Yang, W., Yang, Z.-T., Lu, S.-J., and Liu, J.-X. (2020). “A membrane-associated NAC transcription factor osNTL3 is involved in thermotolerance in rice. Plant Biotechnol. J. 18, 1317–1295. doi: 10.1111/pbi.13297

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, J., Sun, X., Xu, F., Zhang, Y., Zhang, Q., Miao, R., et al. (2018). Suppression of osMDHAR4 enhances heat tolerance by mediating H2O2-induced stomatal closure in rice plants. Rice 11, 38. doi: 10.1186/s12284-018-0230-5

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, Q., Yang, T., Yu, T., Zhang, S., Mao, X., Zhao, J., et al. (2017). Integrating Small RNA Sequencing with QTL Mapping for Identification of miRNAs and Their Target Genes Associated with Heat Tolerance at the Flowering Stage in Rice. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.00043

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, W., Yin, T., Zhao, Y., Wang, X., Wang, K., Shen, Y., et al. (2021). Effects of high temperature on rice grain development and quality formation based on proteomics comparative analysis under field warming. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.746180

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, H., Zeng, B., Zhao, J., Yan, S., Wan, J., and Cao, Z. (2023). “Genetic research progress: heat tolerance in rice. Int. J. Mol. Sci. 24, 85. doi: 10.3390/ijms24087140

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, E., Zeng, S., Zhu, S., Liu, Y., Wu, G., Zhao, K., et al. (2019). Favorable alleles of GRAIN-FILLING RATE1 increase the grain-filling rate and yield of rice1[OPEN. Plant Physiol. 181, 1207–1222. doi: 10.1104/pp.19.00413

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, G., Zha, Z., Cai, H., Dandan, Q., Jia, H., Liu, C., et al. (2020). Dynamic transcriptome analysis of anther response to heat stress during anthesis in thermotolerant rice (Oryza sativa L.). Int. J. Mol. Sci. 21, 1155. doi: 10.3390/ijms21031155

PubMed Abstract | Crossref Full Text | Google Scholar

Liu, J., Zhang, C., Wei, C., Liu, X., Wang, M., Yu, F., et al. (2016). The RING finger ubiquitin E3 ligase osHTAS enhances heat tolerance by promoting H2O2-induced stomatal closure in rice. Plant Physiol. 170, 429–443. doi: 10.1104/pp.15.00879

PubMed Abstract | Crossref Full Text | Google Scholar

Lobell, D., Burke, M., Tebaldi, C., Mastrandrea, M., Falcon, W., and Naylor, R. (2008). Prioritizing climate change adaptation needs for food security in 2030. Sci. (New York N.Y.) 319, 607–610. doi: 10.1126/science.1152339

PubMed Abstract | Crossref Full Text | Google Scholar

Locato, V. and Gara, L. D. (2018). “Programmed cell death in plants: an overview. Methods Mol. Biol. (Clifton N.J.) 1743, 1–8. doi: 10.1007/978-1-4939-7668-3_1

PubMed Abstract | Crossref Full Text | Google Scholar

Lu, S.-J., Yang, Z.-T., Sun, L., Sun, L., Song, Z.-T., and Liu, J.-X. (2012). “Conservation of IRE1-Regulated bZIP74 mRNA Unconventional Splicing in Rice (Oryza Sativa L.) Involved in ER Stress Responses. Mol. Plant 5, 504–145. doi: 10.1093/mp/ssr115

PubMed Abstract | Crossref Full Text | Google Scholar

Lv, Y., Shao, G., Qiu, J., Jiao, G., Sheng, Z., Xie, L., et al. (2017). White leaf and panicle 2, encoding a PEP-associated protein, is required for chloroplast biogenesis under heat stress in rice. J. Exp. Bot. 68, 5147–5160. doi: 10.1093/jxb/erx332

PubMed Abstract | Crossref Full Text | Google Scholar

Lyman, N. B., Jagadish, K. S.V., Nalley, L.L., Dixon, B. L., and Siebenmorgen, T. (2013). “Neglecting rice milling yield and quality underestimates economic losses from high-temperature stress. PloS One 8, e721575. doi: 10.1371/journal.pone.0072157

PubMed Abstract | Crossref Full Text | Google Scholar

Maavimani, M., Jebaraj, S., Raveendran, M., Vanniarajan, C., Balakrishnan, K., and Muthamilan, M. (2014). Cellular membrane thermostability is related to rice (Oryza sativa L) yield under heat stress. Int. J. Trop. Agric. 32, 201–208.

Google Scholar

Matsui, T. and Omasa, K. (2002). “Rice (Oryza sativa L.) cultivars tolerant to high temperature at flowering: anther characteristics. Ann. Bot. 89, 683–875. doi: 10.1093/aob/mcf112

PubMed Abstract | Crossref Full Text | Google Scholar

McCormick, S. (2018). Remembrance of stresses past: heat shock factors and histone hypermethylation are key. Plant Journal: For Cell Mol. Biol. 95, 399–400. doi: 10.1111/tpj.14001

PubMed Abstract | Crossref Full Text | Google Scholar

Miao, C., Xiao, L., Hua, K., Zou, C., Zhao, Y., Bressan, R. A., et al. (2018). Mutations in a subfamily of abscisic acid receptor genes promote rice growth and productivity. Proc. Natl. Acad. Sci. United States America 115, 6058–6063. doi: 10.1073/pnas.1804774115

PubMed Abstract | Crossref Full Text | Google Scholar

Mishra, R. C. and Grover, A. (2016). “ClpB/hsp100 proteins and heat stress tolerance in plants. Crit. Rev. Biotechnol. 36, 862–745. doi: 10.3109/07388551.2015.1051942

PubMed Abstract | Crossref Full Text | Google Scholar

Mishra, A., Singh, B. B., Shakil, N. A., Shamim, M. D., Homa, F., Chaudhary, R., et al. (2024). Effect of high temperature stress on metabolome and aroma in rice grains. Plant Gene 38, 100450. doi: 10.1016/j.plgene.2024.100450

Crossref Full Text | Google Scholar

Mittal, D., Chakrabarti, S., Sarkar, A., Singh, A., and Grover, A. (2009). “Heat shock factor gene family in rice: genomic organization and transcript expression profiling in response to high temperature, low temperature and oxidative stresses. Plant Physiol. Biochem. 47, 785–955. doi: 10.1016/j.plaphy.2009.05.003

PubMed Abstract | Crossref Full Text | Google Scholar

Mittler, R. (2017). ROS are good. Trends Plant Sci. 22, 11–19. doi: 10.1016/j.tplants.2016.08.002

PubMed Abstract | Crossref Full Text | Google Scholar

Mittler, R., Finka, A., and Goloubinoff, P. (2012). “How do plants feel the heat? Trends Biochem. Sci. 37, 118–255. doi: 10.1016/j.tibs.2011.11.007

PubMed Abstract | Crossref Full Text | Google Scholar

Miyahara, K., Wada, T., Sonoda, J., Tsukaguchi, T., Miyazaki, M., Tsubone, M., et al. (2017). Detection and validation of QTLs for milky-white grains caused by high temperature during the ripening period in japonica rice. Breed. Sci. 67, 333–339. doi: 10.1270/jsbbs.16203

PubMed Abstract | Crossref Full Text | Google Scholar

Miyazaki, M., Araki, M., Okamura, K., Ishibashi, Y., Yuasa, T., and Iwaya-Inoue, M. (2013). “Assimilate translocation and expression of sucrose transporter, osSUT1, contribute to high-performance ripening under heat stress in the heat-tolerant rice cultivar genkitsukushi. J. Plant Physiol. 170, 1579–1845. doi: 10.1016/j.jplph.2013.06.011

PubMed Abstract | Crossref Full Text | Google Scholar

Mo, Y., Li, G., Liu, L., Zhang, Y., Li, J., Yang, M., et al. (2023). OsGRF4AA compromises heat tolerance of developing pollen grains in rice. Front. Plant Sci. 14. doi: 10.3389/fpls.2023.1121852

PubMed Abstract | Crossref Full Text | Google Scholar

Mohammed, A. R. and Tarpley, L. (2011). Effects of night temperature, spikelet position and salicylic acid on yield and yield-related parameters of rice (Oryza sativa L.) plants. J. Agron. Crop Sci. 197, 40–49. doi: 10.1111/j.1439-037X.2010.00439.x

Crossref Full Text | Google Scholar

Munir, R., Jan, M., Muhammad, S., Afzal, M., Jan, N., Yasin, M. U., et al. (2023). Detrimental effects of cd and temperature on rice and functions of microbial community in paddy soils. Environ. pollut. 324, 121371. doi: 10.1016/j.envpol.2023.121371

PubMed Abstract | Crossref Full Text | Google Scholar

Murakami, T., Matsuba, S., Funatsuki, H., Kawaguchi, K., Saruyama, H., Tanida, M., et al. (2004). Over-expression of a small heat shock protein, sHSP17.7, confers both heat tolerance and UV-B resistance to rice plants. Mol. Breed. 13, 165–175. doi: 10.1023/B:MOLB.0000018764.30795.c1

Crossref Full Text | Google Scholar

Murata, K., Iyama, Y., Yamaguchi, T., Ozaki, H., Kidani, Y., and Ebitani, T. (2014). “Identification of a novel gene (Apq1) from the indica rice cultivar ‘Habataki’ That improves the quality of grains produced under high temperature stress. Breed. Sci. 64, 273–815. doi: 10.1270/jsbbs.64.273

PubMed Abstract | Crossref Full Text | Google Scholar

Nadarajah, K., Hamid, N. W. A., and Abdul Rahman, N. S. N. (2021). “SA-mediated regulation and control of abiotic stress tolerance in rice. Int. J. Mol. Sci. 22, 55915. doi: 10.3390/ijms22115591

PubMed Abstract | Crossref Full Text | Google Scholar

Nazir, F., Jahan, B., Kumari, S., Iqbal, N., Albaqami, M., Sofo, A., et al. (2023). Brassinosteroid modulates ethylene synthesis and antioxidant metabolism to protect rice (Oryza sativa) against heat stress-induced inhibition of source–sink capacity and photosynthetic and growth attributes. J. Plant Physiol. 289, 154096. doi: 10.1016/j.jplph.2023.154096

PubMed Abstract | Crossref Full Text | Google Scholar

Nevame, A. Y. M., Emon, R. M., Malek, M. A., Hasan, M. M., Alam, M. A., Muharam, F. M., et al. (2018). Relationship between high temperature and formation of chalkiness and their effects on quality of rice. BioMed. Res. Int. 2018, 1653721. doi: 10.1155/2018/1653721

PubMed Abstract | Crossref Full Text | Google Scholar

Nguyen, T., Shen, S., Cheng, M., and Chen, Q. (2022). “Identification of QTLs for heat tolerance at the flowering stage using chromosome segment substitution lines in rice. Genes 13, 22485. doi: 10.3390/genes13122248

PubMed Abstract | Crossref Full Text | Google Scholar

Niu, Y. and Xiang, Y. (2018). An overview of biomembrane functions in plant responses to high-temperature stress. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.00915

PubMed Abstract | Crossref Full Text | Google Scholar

NOAA (2024). 2023 was the world’s warmest year on record, by far | National oceanic and atmospheric administration. Available online at: https://www.noaa.gov/news/2023-was-worlds-warmest-year-on-record-by-far (Accessed September 28, 2025).

Google Scholar

NOAA Climate (2025). Gove. Available online at: https://www.climate.gov/news-features/understanding-climate/climate-change-global-temperature (Accessed June 09, 2025).

Google Scholar

NOAA Global Monitoring Laboratory Trends in CO2 - NOAA global monitoring laboratory. Available online at: https://gml.noaa.gov/ccgg/trends/global.html (Accessed April 28, 2025).

Google Scholar

Noman, A., Aqeel, M., and He, S. (2016). CRISPR-cas9: tool for qualitative and quantitative plant genome editing. Front. Plant Sci. 7. doi: 10.3389/fpls.2016.01740

PubMed Abstract | Crossref Full Text | Google Scholar

Nubankoh, P., Wanchana, S., Saensuk, C., Ruanjaichon, V., Cheabu, S., Vanavichit, A., et al. (2020). QTL-seq reveals genomic regions associated with spikelet fertility in response to a high temperature in rice (Oryza sativa L.). Plant Cell Rep. 39, 149–162. doi: 10.1007/s00299-019-02477-z

PubMed Abstract | Crossref Full Text | Google Scholar

Ogawa, A., Yoshino, S., and Toyofuku, K. (2025). “Metabolomic profiling of heat tolerance during grain filling in rice: comparative analyses of panicles and roots in ‘Fusaotome’ and ‘Akitakomachi.’. Agriculture 15, 22555. doi: 10.3390/agriculture15212255

Crossref Full Text | Google Scholar

Ohama, N., Sato, H., Shinozaki, K., and Yamaguchi-Shinozaki, K. (2017). “Transcriptional regulatory network of plant heat stress response. Trends Plant Sci. 22, 53–655. doi: 10.1016/j.tplants.2016.08.015

PubMed Abstract | Crossref Full Text | Google Scholar

Our World in Data. Greenhouse gas emissions. Available online at: https://ourworldindata.org/grapher/total-ghg-emissions (Accessed April 19, 2025).

Google Scholar

Pan, Y. H., Chen, L., Zhu, X. Y., Li, J. C., Rashid, M. A. R. R., Chen, C., et al. (2023). Utilization of natural alleles for heat adaptability QTLs at the flowering stage in rice. BMC Plant Biol. 23, 256. doi: 10.1186/s12870-023-04260-5

PubMed Abstract | Crossref Full Text | Google Scholar

Pandian, S., Rakkammal, K., Rency, A., Muthuramalingam, P., Pandian, S., and Ramesh, M. (2020). Abiotic stress and applications of omics approaches to develop stress tolerance in agronomic crops. Agronomic CropsAgronomic Crops 557–578. doi: 10.1007/978-981-15-0025-1_26

Crossref Full Text | Google Scholar

Pang, Y., Hu, Y., and Bao, J. (2021). “Comparative phosphoproteomic analysis reveals the response of starch metabolism to high-temperature stress in rice endosperm. Int. J. Mol. Sci. 22, 105465. doi: 10.3390/ijms221910546

PubMed Abstract | Crossref Full Text | Google Scholar

Park, J.-R., Kim, E.-G., Jang, Y.-H., and Kim, K.-M. (2021). “Screening and identification of genes affecting grain quality and spikelet fertility during high-temperature treatment in grain filling stage of rice. BMC Plant Biol. 21, 2635. doi: 10.1186/s12870-021-03056-9

PubMed Abstract | Crossref Full Text | Google Scholar

Park, S. I., Kwon, H. J., Cho, M. H., Song, J. S., Kim, B. G., Baek, J., et al. (2021). The osERF115/AP2EREBP110 transcription factor is involved in the multiple stress tolerance to heat and drought in rice plants. Int. J. Mol. Sci. 22, 13. doi: 10.3390/ijms22137181

PubMed Abstract | Crossref Full Text | Google Scholar

Park, J.-R., Yang, W.-T., Kim, D.-H., and Kim, K.-M. (2020). “Identification of a novel gene, osbht, in response to high temperature tolerance at booting stage in rice. Int. J. Mol. Sci. 21, 16. doi: 10.3390/ijms21165862

PubMed Abstract | Crossref Full Text | Google Scholar

Payman, S. H., Bakhshipour, A., and Zareiforoush, H. (2018). Development of an expert vision-based system for inspecting rice quality indices. Qual. Assur. Saf. Crops Foods 10, 103–114. doi: 10.3920/QAS2017.1109

Crossref Full Text | Google Scholar

Peng, S., Huang, J., Sheehy, J. E., Laza, R. C., Visperas, R. M., Zhong, X., et al. (2004). Rice yields decline with higher night temperature from global warming. Proc. Natl. Acad. Sci. United States America 101, 9971–9975. doi: 10.1073/pnas.0403720101

PubMed Abstract | Crossref Full Text | Google Scholar

Perdomo, J. A., Capó-Bauçà, S., Carmo-Silva, E., and Galmés, J. (2017). “Rubisco and rubisco activase play an important role in the biochemical limitations of photosynthesis in rice, wheat, and maize under high temperature and water deficit. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.00490

PubMed Abstract | Crossref Full Text | Google Scholar

Prasanth, V. V., Babu, M. S., Basava, R. K., Venkata, V. G. N. T., Mangrauthia, S. K., Voleti, S. R., et al. (2017). Trait and marker associations in oryza nivara and O. Rufipogon derived rice lines under two different heat stress conditions. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01819

PubMed Abstract | Crossref Full Text | Google Scholar

Prerostova, S., Jarosova, J., Dobrev, P. I., Hluskova, L., Motyka, V., Filepova, R., et al. (2022). Heat stress targeting individual organs reveals the central role of roots and crowns in rice stress responses. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.799249

PubMed Abstract | Crossref Full Text | Google Scholar

PS, S., Mithra, A. S. V., Prakash, C., Ramkumar, M. K., Tiwari, R., Mohapatra, T., et al. (2017). High resolution mapping of QTLs for heat tolerance in rice using a 5K SNP array. Rice 10, 28. doi: 10.1186/s12284-017-0167-0

PubMed Abstract | Crossref Full Text | Google Scholar

Qi, Y., Wang, H., Zou, Y., Liu, C., Liu, Y., Wang, Y., et al. (2011). Over-expression of mitochondrial heat shock protein 70 suppresses programmed cell death in rice. FEBS Lett. 585, 231–239. doi: 10.1016/j.febslet.2010.11.051

PubMed Abstract | Crossref Full Text | Google Scholar

Qiao, B., Zhang, Q., Liu, D., Wang, H., Yin, J., Wang, R., et al. (2015). A calcium-binding protein, rice annexin osANN1, enhances heat stress tolerance by modulating the production of H2O2. J. Exp. Bot. 66, 5853–5866. doi: 10.1093/jxb/erv294

PubMed Abstract | Crossref Full Text | Google Scholar

Qin, Q., Zhao, Y., Zhang, J., Chen, L., Si, W., and Jiang, H. (2022). “A maize heat shock factor zmHsf11 negatively regulates heat stress tolerance in transgenic plants. BMC Plant Biol. 22, 4065. doi: 10.1186/s12870-022-03789-1

PubMed Abstract | Crossref Full Text | Google Scholar

Qiu, Z., Kang, S., He, L., Zhao, J., Zhang, S., Hu, J., et al. (2018). The newly identified heat-stress sensitive albino 1 gene affects chloroplast development in rice. Plant Science: Int. J. Exp. Plant Biol. 267, 168–179. doi: 10.1016/j.plantsci.2017.11.015

PubMed Abstract | Crossref Full Text | Google Scholar

Qiu, R., Yao, P., Yang, J., Hou, J., Xiao, H., Wu, Y., et al. (2025). OsIAA7 enhances heat stress tolerance by inhibiting the activity of osARF6 in rice. Int. J. Biol. Macromolecules 288, 138746. doi: 10.1016/j.ijbiomac.2024.138746

PubMed Abstract | Crossref Full Text | Google Scholar

Qiu, F., Zheng, Y., Lin, Y., Woldegiorgis, S. T., Xu, S., Feng, C., et al. (2023). Integrated ATAC-seq and RNA-seq data analysis to reveal osbZIP14 function in rice in response to heat stress. Int. J. Mol. Sci. 24, 5619. doi: 10.3390/ijms24065619

PubMed Abstract | Crossref Full Text | Google Scholar

Qiu, Z., Zhu, L., He, L., Chen, D., Zeng, D., Chen, G., et al. (2019). DNA damage and reactive oxygen species cause cell death in the rice local lesions 1 mutant under high light and high temperature. New Phytol. 222, 349–365. doi: 10.1111/nph.15597

PubMed Abstract | Crossref Full Text | Google Scholar

Qu, Y., Sakoda, K., Fukayama, H., Kondo, E., Suzuki, Y., Makino, A., et al. (2021). Overexpression of both rubisco and rubisco activase rescues rice photosynthesis and biomass under heat stress. Plant Cell Environ. 44, 2308–2320. doi: 10.1111/pce.14051

PubMed Abstract | Crossref Full Text | Google Scholar

Radha, B., Sunitha, N. C., Sah, R. P., Azharudheen, M. T. P., Krishna, G. K., Umesh, D. K., et al. (2022). Physiological and molecular implications of multiple abiotic stresses on yield and quality of rice. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.996514

PubMed Abstract | Crossref Full Text | Google Scholar

Rang, Z. W., Jagadish, S. V. K., Zhou, Q. M., Craufurd, P. Q., and Heuer, S. (2011). Effect of high temperature and water stress on pollen germination and spikelet fertility in rice. Environ. Exp. Bot. 70, 58–65. doi: 10.1016/j.envexpbot.2010.08.009

Crossref Full Text | Google Scholar

Rangappa, K., Choudhury, B. U., Kumar, A., Das, S. P., Ayam, G., Hazarika, S., et al. (2024). Comparative stress physiological analysis of indigenous rice cultivars of eastern himalayan region under elevated temperature of changing climate. Plant Physiol. Rep. 535–551. doi: 10.1007/s40502-024-00796-2

Crossref Full Text | Google Scholar

Ravikiran, K. T., Krishnan, S. G., Abhijith, K. P., Bollinedi, H., Nagarajan, M., Vinod, K. K., et al. (2022). Genome-wide association mapping reveals novel putative gene candidates governing reproductive stage heat stress tolerance in rice. Front. Genet. 13. doi: 10.3389/fgene.2022.876522

PubMed Abstract | Crossref Full Text | Google Scholar

Raza, Q., Riaz, A., Bashir, K., and Sabar, M. (2020). Reproductive tissues-specific meta-QTLs and candidate genes for development of heat-tolerant rice cultivars. Plant Mol. Biol. 104, 97–112. doi: 10.1007/s11103-020-01027-6

PubMed Abstract | Crossref Full Text | Google Scholar

Ren, H., Bao, J., Gao, Z., Sun, D., Zheng, S., and Bai, J. (2023). How rice adapts to high temperatures. Front. Plant Sci. 14. doi: 10.3389/fpls.2023.1137923

PubMed Abstract | Crossref Full Text | Google Scholar

Ren, D., Ding, C., and Qian, Q. (2023). “Molecular bases of rice grain size and quality for optimized productivity. Sci. Bull. 68, 314–505. doi: 10.1016/j.scib.2023.01.026

PubMed Abstract | Crossref Full Text | Google Scholar

Ren, Y., Huang, Z., Jiang, H., Wang, Z., Wu, F., Xiong, Y., et al. (2021). A heat stress responsive NAC transcription factor heterodimer plays key roles in rice grain filling. J. Exp. Bot. 72, 2947–2964. doi: 10.1093/jxb/erab027

PubMed Abstract | Crossref Full Text | Google Scholar

Rerksiri, W., Zhang, X., Xiong, H., and Chen, X. (2013). Expression and promoter analysis of six heat stress-inducible genes in rice. Sci. World J. 2013, 397401. doi: 10.1155/2013/397401

PubMed Abstract | Crossref Full Text | Google Scholar

Restrepo-Diaz, H. and Garces, G. (2013). Response of Rice Plants to Heat Stress during Initiation of Panicle Primordia or Grain-Filling Phases Response of Rice Plants to Heat Stress during Initiation of Panicle Primordia or Grain-Filling Phases. J. Stress Physiol. Biochem. 9, 318–325.

Google Scholar

Rezaul, I. M., Baohua, F., Tingting, C., Weimeng, F., Caixia, Z., Longxing, T., et al. (2019). Abscisic acid prevents pollen abortion under high-temperature stress by mediating sugar metabolism in rice spikelets. Physiologia Plantarum 165, 644–663. doi: 10.1111/ppl.12759

PubMed Abstract | Crossref Full Text | Google Scholar

Rizhsky, L., Liang, H., and Mittler, R. (2002). “The combined effect of drought stress and heat shock on gene expression in tobacco. Plant Physiol. 130, 1143–1515. doi: 10.1104/pp.006858

PubMed Abstract | Crossref Full Text | Google Scholar

Rizhsky, L., Liang, H., Shuman, J., Shulaev, V., Davletova, S., and Mittler, R. (2004). “When defense pathways collide. The response of arabidopsis to a combination of drought and heat stress. Plant Physiol. 134, 1683–1965. doi: 10.1104/pp.103.033431

PubMed Abstract | Crossref Full Text | Google Scholar

Robson, J. K., Ferguson, J. N., McAusland, L., Tranchant-Dubreuil, C., Cubry, P., Sabot, F., et al. (2023). Chlorophyll fluorescence-based high-throughput phenotyping facilitates the genetic dissection of photosynthetic heat tolerance in african (Oryza glaberrima) and asian (Oryza sativa) rice. J. Exp. Bot. 74, 5181–5197. doi: 10.1093/jxb/erad239

PubMed Abstract | Crossref Full Text | Google Scholar

Saidi, Y., Peter, M., Finka, A., Cicekli, C., Vigh, L., and Goloubinoff, P. (2010). “Membrane lipid composition affects plant heat sensing and modulates ca2+-dependent heat shock response. Plant Signaling Behav. 5, 1530–1335. doi: 10.4161/psb.5.12.13163

PubMed Abstract | Crossref Full Text | Google Scholar

Sailaja, B., Subrahmanyam, D., Neelamraju, S., Vishnukiran, T., Rao, Y. V., Vijayalakshmi, P., et al. (2015). Integrated physiological, biochemical, and molecular analysis identifies important traits and mechanisms associated with differential response of rice genotypes to elevated temperature. Front. Plant Sci. 6. doi: 10.3389/fpls.2015.01044

PubMed Abstract | Crossref Full Text | Google Scholar

Sailaja, B., Voleti, S. R., Subrahmanyam, D., Sarla, N., Prasanth, V. V., Bhadana, V. P., et al. (2014). Prediction and expression analysis of miRNAs associated with heat stress in oryza sativa. Rice Sci. 21, 3–12. doi: 10.1016/S1672-6308(13)60164-X

Crossref Full Text | Google Scholar

Sánchez, B., Rasmussen, A., and Porter, J. R. (2014). “Temperatures and the growth and development of maize and rice: A review. Global Change Biol. 20, 408–175. doi: 10.1111/gcb.12389

PubMed Abstract | Crossref Full Text | Google Scholar

Sandhu, J., Irvin, L., Liu, K., Staswick, P., Zhang, C., and Walia, H. (2021). “Endoplasmic reticulum stress pathway mediates the early heat stress response of developing rice seeds. Plant Cell Environ. 44, 2604–2245. doi: 10.1111/pce.14103

PubMed Abstract | Crossref Full Text | Google Scholar

Sangwan, V., Orvar, B. L., Beyerly, J., Hirt, H., and Dhindsa, R. S. (2002). “Opposite changes in membrane fluidity mimic cold and heat stress activation of distinct plant MAP kinase pathways. Plant Journal: For Cell Mol. Biol. 31, 629–385. doi: 10.1046/j.1365-313x.2002.01384.x

PubMed Abstract | Crossref Full Text | Google Scholar

Sarangthem, K., Yumlembam, S., Benazir, S., Yendrembam, R., and Mikawlrawng, K. (2021). Current understanding of the mechanisms of heat stress tolerance in rice (Oryza Sativa L.). J. Exp. Biol. Agric. Sci. 9, S321–S329. doi: 10.18006/2021.9(Spl-3-NRMCSSA_2021).S321.S329

Crossref Full Text | Google Scholar

Sarkar, N. K., Kotak, S., Agarwal, M., Kim, Y.-K., and Grover, A. (2019). “Silencing of class I small heat shock proteins affects seed-related attributes and thermotolerance in rice seedlings. Planta 251, 265. doi: 10.1007/s00425-019-03318-9

PubMed Abstract | Crossref Full Text | Google Scholar

Sarker, N. K., Kotak, S., Agarwal, M., Kim, Y. K., Grover, A., et al. (2025). Hot off the genes: uncovering differentially expressed genes in Malaysian weedy rice in response to heat stress. Euphytica 221, 115. doi: 10.1007/s10681-025-03569-6

Crossref Full Text | Google Scholar

Sato, H., Todaka, D., Kudo, M., Mizoi, J., Kidokoro, S., Zhao, Y., et al. (2016). The arabidopsis transcriptional regulator DPB3–1 enhances heat stress tolerance without growth retardation in rice. Plant Biotechnol. J. 14, 1756–1767. doi: 10.1111/pbi.12535

PubMed Abstract | Crossref Full Text | Google Scholar

Scafaro, A. P., Atwell, B. J., Muylaert, S., Van Reusel, B., Ruiz, G. A., Van Rie, J., et al. (2018). A thermotolerant variant of rubisco activase from a wild relative improves growth and seed yield in rice under heat stress. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01663

PubMed Abstract | Crossref Full Text | Google Scholar

Schaarschmidt, S., Lawas, L. M. F., Glaubitz, U., Li, X., Erban, A., Kopka, J., et al. (2020). Season affects yield and metabolic profiles of rice (Oryza sativa) under high night temperature stress in the field. Int. J. Mol. Sci. 21, 3187. doi: 10.3390/ijms21093187

PubMed Abstract | Crossref Full Text | Google Scholar

Seem, K., Ghosh, A., Kumar, S., and Mohapatra, T. (2024). Epigenomics of adaptive plasticity of rice under changing climatic conditions grown by direct-sowing as well as transplanting. Arch. Mol. Biol. Genet. 3, 5–19. doi: 10.33696/genetics.3.015

Crossref Full Text | Google Scholar

Seo, J., Lee, S.-M., Han, J.-H., Shin, N.-H., Koh, H.-J., and Chin, J. H. (2019). Identification of yield and yield-related quantitative trait loci for the field high temperature condition in backcross populations of rice ( Oryza sativa L.). Plant Breed. Biotechnol. 7, 415–426. doi: 10.9787/PBB.2019.7.4.415

Crossref Full Text | Google Scholar

Shanmugavadivel, P. S., Prakash, C., and Mithra, S.V.A. (2019). “Chapter 42 - molecular approaches for dissecting and improving drought and heat tolerance in rice,” in Advances in rice research for abiotic stress tolerance. Eds. Hasanuzzaman, M., Fujita, M., Nahar, K., and Biswas, J. K. (Swaston, Cambridge: Woodhead Publishing). doi: 10.1016/B978-0-12-814332-2.00042-3

Crossref Full Text | Google Scholar

Sharma, E., Borah, P., Kaur, A., Bhatnagar, A., Mohapatra, T., Kapoor, S., et al. (2021). A comprehensive transcriptome analysis of contrasting rice cultivars highlights the role of auxin and ABA responsive genes in heat stress response. Genomics 113, 1247–1261. doi: 10.1016/j.ygeno.2021.03.007

PubMed Abstract | Crossref Full Text | Google Scholar

Sharma, N., Singh, B., Krishnan, S. G., Bollinedi, H., Mandal, P. K., Lal, M. K., et al. (2024). Higher grain filling rate in inferior spikelets of tolerant rice genotype offset grain yield loss under post-anthesis high night temperature. Rice Sci. 572–586. doi: 10.1016/j.rsci.2024.06.003

Crossref Full Text | Google Scholar

Sharma, D., Singh, R., Tiwari, R., Kumar, R., and Gupta, V. K. (2019). Wheat responses and tolerance to terminal heat stress: A review, in Wheat production in changing environments: responses, adaptation and tolerance. Eds. Hasanuzzaman, M., Nahar, K., and Hossain, A. (Gateway East, Singapore: Springer). doi: 10.1007/978-981-13-6883-7_7

Crossref Full Text | Google Scholar

Shekhawat, K., Almeida-Trapp, M., García-Ramírez, G. X., and Hirt, H. (2022). “Beat the heat: plant- and microbe-mediated strategies for crop thermotolerance. Trends Plant Sci. 27, 802–135. doi: 10.1016/j.tplants.2022.02.008

PubMed Abstract | Crossref Full Text | Google Scholar

Shen, H., Zhong, X., Zhao, F., Wang, Y., Yan, B., Li, Q., et al. (2015). Overexpression of receptor-like kinase ERECTA improves thermotolerance in rice and tomato. Nat. Biotechnol. 33, 996–1003. doi: 10.1038/nbt.3321

PubMed Abstract | Crossref Full Text | Google Scholar

Shi, W., Muthurajan, R., Rahman, H., Selvam, J., Peng, S., Zou, Y., et al. (2013). Source-sink dynamics and proteomic reprogramming under elevated night temperature and their impact on rice yield and grain quality. New Phytol. 197, 825–837. doi: 10.1111/nph.12088

PubMed Abstract | Crossref Full Text | Google Scholar

Shi, W., Yang, J., Kumar, R., Zhang, X., Impa, S. M., Xiao, G., et al. (2022). Heat stress during gametogenesis irreversibly damages female reproductive organ in rice. Rice 15, 32. doi: 10.1186/s12284-022-00578-0

PubMed Abstract | Crossref Full Text | Google Scholar

Shi, W., Yin, X., Struik, P. C., Solis, C., Xie, F., Schmidt, R. C., et al. (2017). High day- and night-time temperatures affect grain growth dynamics in contrasting rice genotypes. J. Exp. Bot. 68, 5233–5245. doi: 10.1093/jxb/erx344

PubMed Abstract | Crossref Full Text | Google Scholar

Shiraya, T., Mori, T., Maruyama, T., Sasaki, M., Takamatsu, T., Oikawa, K., et al. (2015). Golgi/plastid-type manganese superoxide dismutase involved in heat-stress tolerance during grain filling of rice. Plant Biotechnol. J. 13, 1251–1263. doi: 10.1111/pbi.12314

PubMed Abstract | Crossref Full Text | Google Scholar

Shirdelmoghanloo, H., Chen, K., Paynter, B. H., Angessa, T. T., Westcott, S., Khan, H. A., et al. (2022). Grain-filling rate improves physical grain quality in barley under heat stress conditions during the grain-filling period. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.858652

PubMed Abstract | Crossref Full Text | Google Scholar

Shrestha, S., Mahat, J., Shrestha, J., K.c., M., and Paudel, K. (2022). “Influence of high-temperature stress on rice growth and development. A review. Heliyon 8, e126515. doi: 10.1016/j.heliyon.2022.e12651

PubMed Abstract | Crossref Full Text | Google Scholar

Singh, A., Kumar, M., Chakdar, H., Pandiyan, K., Kumar, S. C., Zeyad, M. T., et al. (2022). Influence of host genotype in establishing root associated microbiome of indica rice cultivars for plant growth promotion. Front. Microbiol. 13. doi: 10.3389/fmicb.2022.1033158

PubMed Abstract | Crossref Full Text | Google Scholar

Singh, A., Mazahar, S., Chapadgaonkar, S. S., Giri, P., and Shourie, A. (2023). Phyto-microbiome to mitigate abiotic stress in crop plants. Front. Microbiol. 14. doi: 10.3389/fmicb.2023.1210890

PubMed Abstract | Crossref Full Text | Google Scholar

Singh, S., Praveen, A., Dudha, N., and Bhadrecha, P. (2024). “Integrating physiological and multi-omics methods to elucidate heat stress tolerance for sustainable rice production. Physiol. Mol. Biol. Plants. 1185–1208. doi: 10.1007/s12298-024-01480-3

PubMed Abstract | Crossref Full Text | Google Scholar

Singha, D. L., Maharana, J., Panda, D., Dehury, B., Modi, M. K., and Singh, S. (2021). “Understanding the Thermal Response of Rice Eukaryotic Transcription Factor eIF4A1 towards Dynamic Temperature Stress: Insights from Expression Profiling and Molecular Dynamics Simulation. J. Biomolecular Structure Dynamics 39, 2575–2845. doi: 10.1080/07391102.2020.1751295

PubMed Abstract | Crossref Full Text | Google Scholar

Sita, K., Sehgal, A., HanumanthaRao, B., Nair, R. M., Prasad, P. V. V., Kumar, S., et al. (2017). Food legumes and rising temperatures: effects, adaptive functional mechanisms specific to reproductive growth stage and strategies to improve heat tolerance. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01658

PubMed Abstract | Crossref Full Text | Google Scholar

Soda, N., Gupta, B. K., Anwar, K., Sharan, P., Govindjee., Singla-Pareek, S. L., et al. (2018). Rice intermediate filament, osIF, stabilizes photosynthetic machinery and yield under salinity and heat stress. Sci. Rep. 8, 4072. doi: 10.1038/s41598-018-22131-0

PubMed Abstract | Crossref Full Text | Google Scholar

Song, Y., Chen, Q., Ci, D., Shao, X., and Zhang, D. (2014). Effects of high temperature on photosynthesis and related gene expression in poplar. BMC Plant Biol. 14, 111. doi: 10.1186/1471-2229-14-111

PubMed Abstract | Crossref Full Text | Google Scholar

Sreenivasulu, N., Butardo, V. M., Misra, G., Cuevas, R. P., Anacleto, R., and Kavi Kishor, P. B. (2015). “Designing climate-resilient rice with ideal grain quality suited for high-temperature stress. J. Exp. Bot. 66, 1737–1485. doi: 10.1093/jxb/eru544

PubMed Abstract | Crossref Full Text | Google Scholar

Stephen, K., Beena, R., Neethu, M., and Shanija, S. (2022). “Identification of heat-tolerant rice genotypes and their molecular characterisation using SSR markers. Plant Sci. Today. 802–813. doi: 10.14719/pst.1639

Crossref Full Text | Google Scholar

Sun, J.-L., Li, J.-Y., Wang, M.-J., Song, Z.-T., and Liu, J.-X. (2021). “Protein quality control in plant organelles: current progress and future perspectives. Mol. Plant 14, 95–1145. doi: 10.1016/j.molp.2020.10.011

PubMed Abstract | Crossref Full Text | Google Scholar

Surender, R., Lavanya, K., Fiyaz, A., Chiranjeevi, M., Shiva Prasad, G., and Rao, L. V. S. (2021). “Screening of rice (O. Sativa L.) genotypes for heat tolerance to develop climate resilient rice genotypes. Pharma Innovation J. 10, 132–435.

Google Scholar

Suriyasak, C., Harano, K., Tanamachi, K., Matsuo, K., Tamada, A., Iwaya-Inoue, M., et al. (2017). Reactive oxygen species induced by heat stress during grain filling of rice (Oryza sativa L.) are involved in occurrence of grain chalkiness. J. Plant Physiol. 216, 52–57. doi: 10.1016/j.jplph.2017.05.015

PubMed Abstract | Crossref Full Text | Google Scholar

Suriyasak, C., Kawaguchi, R., Matsumoto, R., Sawada, Y., Nong, H. T., Hamaoka, N., et al. (2025). Adaptive memory induced by heat stress during grain filling enhances subsequent thermotolerance in rice (Oryza sativa L.). Sci. Rep. 15, 14135. doi: 10.1038/s41598-025-99146-x

PubMed Abstract | Crossref Full Text | Google Scholar

Suriyasak, C., Oyama, Y., Ishida, T., Mashiguchi, K., Yamaguchi, S., Hamaoka, N., et al. (2020). Mechanism of delayed seed germination caused by high temperature during grain filling in rice (Oryza sativa L.). Sci. Rep. 10, 17378. doi: 10.1038/s41598-020-74281-9

PubMed Abstract | Crossref Full Text | Google Scholar

Szymańska, R., Ślesak, I., Orzechowska, A., and Kruk, J. (2017). Physiological and biochemical responses to high light and temperature stress in plants. Environ. Exp. Bot. 139, 165–177. doi: 10.1016/j.envexpbot.2017.05.002

Crossref Full Text | Google Scholar

Tabassum, R., Dosaka, T., Ichida, H., Morita, R., Ding, Y., Abe, T., et al. (2020). FLOURY ENDOSPERM11–2 encodes plastid HSP70–2 involved with the temperature-dependent chalkiness of rice (Oryza sativa L.) grains. Plant J. 103, 604–616. doi: 10.1111/tpj.14752

PubMed Abstract | Crossref Full Text | Google Scholar

Takehara, K., Murata, K., Yamaguchi, T., Yamaguchi, K., Chaya, G., Kido, S., et al. (2018). Thermo-responsive allele of sucrose synthase 3 (Sus3) provides high-temperature tolerance during the ripening stage in rice (Oryza sativa L.). Breed. Sci. 68, 336–342. doi: 10.1270/jsbbs.18007

PubMed Abstract | Crossref Full Text | Google Scholar

Tang, Y., Gao, C. C., Gao, Y., Yang, Y., Shi, B., Yu, J. L., et al. (2020). OsNSUN2-mediated 5-methylcytosine mRNA modification enhances rice adaptation to high temperature. Dev. Cell 53, 272–286.e7. doi: 10.1016/j.devcel.2020.03.009

PubMed Abstract | Crossref Full Text | Google Scholar

Tang, R.-S., Zheng, J.-C., Jin, Z.-Q., Zhang, D.-D., Huang, Y.-H., and Chen, L.-G. (2008). “Possible correlation between high temperature-induced floret sterility and endogenous levels of IAA, GAs and ABA in rice (Oryza sativa L.). Plant Growth Regul. 54, 37–435. doi: 10.1007/s10725-007-9225-8

Crossref Full Text | Google Scholar

Tao, L.-X., TAN, H.-J., WANG, X., CAO, L.-Y., Jian, S. O. N. G., and Shi-Hua, CHENG. (2008). Effects of high temperature stress on flowering and grain-setting characteristics for guodao 6. Acta Agronomica Sin. 34, 609–674. doi: 10.1016/S1875-2780(08)60027-9

Crossref Full Text | Google Scholar

Taratima, W., Chuanchumkan, C., Maneerattanarungroj, P., Trunjaruen, A., Theerakulpisut, P., and Dongsansuk, A. (2022). “Effect of heat stress on some physiological and anatomical characteristics of rice (Oryza sativa L.) cv. KDML105 callus and seedling. Biology 11, 115. doi: 10.3390/biology11111587

PubMed Abstract | Crossref Full Text | Google Scholar

Tazib, T., Kobayashi, Y., Koyama, H., and Matsui, T. (2015). “QTL analyses for anther length and dehiscence at flowering as traits for the tolerance of extreme temperatures in rice (Oryza sativa L.). Euphytica 203, 629–425. doi: 10.1007/s10681-014-1291-1

Crossref Full Text | Google Scholar

Tenorio, F. A., Ye, C.-r., Redona, E., Sierra, S., Laza, M., and Argayoso, M. (2013). Screening rice genetic resource for heat tolerance. SABRAO J. Breed. Genet. 45, 341–351.

Google Scholar

Tingting, L., Jinpeng, Z., Xi, Y., Kejian, W., Yuchun, R., and Chun, W. (2023). “Development and application of prime editing in plants. Rice Sci. 30, 509–225. doi: 10.1016/j.rsci.2023.07.005

Crossref Full Text | Google Scholar

Tiwari, S., Prasad, V., Chauhan, P. S., and Lata, C. (2017). Bacillus amyloliquefaciens confers tolerance to various abiotic stresses and modulates plant response to phytohormones through osmoprotection and gene expression regulation in rice. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.01510

PubMed Abstract | Crossref Full Text | Google Scholar

UN DESA World population prospects. Available online at: https://population.un.org/wpp/ (Accessed November 28, 2025).

Google Scholar

USDA (2023). Data and analysis | USDA foreign agricultural service. Available online at: https://fas.usda.gov/data (Accessed October 05, 2025).

Google Scholar

Varshney, R. K., Singh, V. K., Kumar, A., Powell, W., and Sorrells, M. E. (2018). Can genomics deliver climate-change ready crops? Curr. Opin. Plant Biology AGRI 2017 45, 205–211. doi: 10.1016/j.pbi.2018.03.007

PubMed Abstract | Crossref Full Text | Google Scholar

Venuprasad, R., Lafitte, H., and Atlin, G. (2007). Response to direct selection for grain yield under drought stress in rice. Crop Sci. - Crop Sci. 47, 285–293. doi: 10.2135/cropsci2006.03.0181

Crossref Full Text | Google Scholar

Visakh, R. L., Anand, S., Arya, S. N., Sasmita, B., Jha, U. C., Sah, R. P., et al. (2024). Rice heat tolerance breeding: A comprehensive review and forward gaze. Rice Sci. 375–400. doi: 10.1016/j.rsci.2024.02.004

Crossref Full Text | Google Scholar

Vitoriano, C. B. and Calixto, C. P. G. (2021). Reading between the lines: RNA-seq data mining reveals the alternative message of the rice leaf transcriptome in response to heat stress. Plants 10, 85. doi: 10.3390/plants10081647

PubMed Abstract | Crossref Full Text | Google Scholar

Vivitha, P., Raveendran, M., and Vijayalakshmi, D. (2017). Introgression of QTLs controlling spikelet fertility maintains membrane integrity and grain yield in improved white ponni derived progenies exposed to heat stress. Rice Sci. 24, 32–40. doi: 10.1016/j.rsci.2016.05.006

Crossref Full Text | Google Scholar

Wada, H., Hatakeyama, Y., Nakashima, T., Nonami, H., Erra-Balsells, R., Hakata, M., et al. (2020). On-site single pollen metabolomics reveals varietal differences in phosphatidylinositol synthesis under heat stress conditions in rice. Sci. Rep. 10, 2013. doi: 10.1038/s41598-020-58869-9

PubMed Abstract | Crossref Full Text | Google Scholar

Wan, J., Zhang, J., Zan, X., Zhu, J., Chen, H., Li, X., et al. (2023). Overexpression of rice histone H1 gene reduces tolerance to cold and heat stress. Plants (Basel Switzerland) 12, 2408. doi: 10.3390/plants12132408

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, D. R., Bunce, J. A., Tomecek, M. B., Gealy, D., McClung, A., McCouch, S. R., et al. (2016). Evidence for divergence of response in indica, japonica, and wild rice to high CO2 × Temperature interaction. Global Change Biol. 22, 2620–2632. doi: 10.1111/gcb.13279

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, C., Caragea, D., Narayana, N. K., Hein, N. T., Bheemanahalli, R., Somayanda, I. M., et al. (2022). Deep learning based high-throughput phenotyping of chalkiness in rice exposed to high night temperature. Plant Methods 18, 9. doi: 10.1186/s13007-022-00839-5

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, Q.-L., Chen, J.-H., He, N.-Y., and Guo, F.-Q. (2018). “Metabolic reprogramming in chloroplasts under heat stress in plants. Int. J. Mol. Sci. 19, 8495. doi: 10.3390/ijms19030849

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, X., Chen, J., Liu, C., Luo, J., Yan, X., Ai, A., et al. (2019). Over-expression of a protein disulfide isomerase gene from methanothermobacter thermautotrophicus, enhances heat stress tolerance in rice. Gene 684, 124–130. doi: 10.1016/j.gene.2018.10.064

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, G.-F., Li, W.-Q., Li, W.-Y., Wu, G.-L., Zhou, C.-Y., and Chen, K.-M. (2013). “Characterization of rice NADPH oxidase genes and their expression under various environmental conditions. Int. J. Mol. Sci. 14, 9440–9585. doi: 10.3390/ijms14059440

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, D., Li, X.-F., Zhou, Z.-J., Feng, X.-P., Yang, W.-J., and Jiang, D.-A. (2010). “Two rubisco activase isoforms may play different roles in photosynthetic heat acclimation in the rice plant. Physiologia Plantarum 139, 55–675. doi: 10.1111/j.1399-3054.2009.01344.x

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, J., Liu, X. I., Zhang, A. N., Ren, Y., Wu, F., Wang, G., et al. (2019). A cyclic nucleotide-gated channel mediates cytoplasmic calcium elevation and disease resistance in rice. Cell Res. 29, 820–831. doi: 10.1038/s41422-019-0219-7

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, Y., Wang, Y., Liu, X., Zhou, J., Deng, H., Zhang, G., et al. (2022). Dehydration-responsive element binding protein 1C, 1E, and 1G promote stress tolerance to chilling, heat, drought, and salt in rice. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.851731

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, D., Qin, B., Li, X., Tang, D., Zhang, Y., Cheng, Z., et al. (2016). Nucleolar DEAD-box RNA helicase TOGR1 regulates thermotolerant growth as a pre-rRNA chaperone in rice. PloS Genet. 12, e1005844. doi: 10.1371/journal.pgen.1005844

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, Y., Sun, F., Cao, H., Peng, H., Ni, Z., Sun, Q., et al. (2012). TamiR159 directed wheat taGAMYB cleavage and its involvement in anther development and heat response. PloS One 7, e48445. doi: 10.1371/journal.pone.0048445

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, H., Lu, S., Guan, X., Jiang, Y., Wang, B., Hua, J., et al. (2022). WGCNA analysis identifies the hub genes related to heat stress in seedling of rice (Oryza sativa L.). Genes 13, 6. doi: 10.3390/genes13061020

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, J., Xu, J., Wang, L., Zhou, M., Nian, J., Chen, M., et al. (2023). Semi-rolled leaf 10 stabilizes catalase isozyme B to regulate leaf morphology and thermotolerance in rice (Oryza sativa L.). Plant Biotechnol. J. 21, 819–838. doi: 10.1111/pbi.13999

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, Y., Yu, Y., Huang, M., Gao, P., Chen, H., Liu, M., et al. (2020). Transcriptomic and proteomic profiles of II YOU 838 (Oryza sativa) provide insights into heat stress tolerance in hybrid rice. PeerJ 8, e8306. doi: 10.7717/peerj.8306

PubMed Abstract | Crossref Full Text | Google Scholar

Wang, B., Zhong, Z., Wang, X., Han, X., Yu, D., Wang, C., et al. (2020). Knockout of the osNAC006 transcription factor causes drought and heat sensitivity in rice. Int. J. Mol. Sci. 21, 7. doi: 10.3390/ijms21072288

PubMed Abstract | Crossref Full Text | Google Scholar

Waqas, M., Khan, A. L., Shahzad, R., Ullah, I., Khan, A. R., and Lee, I.-J. (2015). “Mutualistic fungal endophytes produce phytohormones and organic acids that promote japonica rice plant growth under prolonged heat stress. J. Zhejiang University. Science. B 16, 10115. doi: 10.1631/jzus.B1500081

PubMed Abstract | Crossref Full Text | Google Scholar

Wei, Z., Yuan, Q., Lin, H., Li, X., Zhang, C., Gao, H., et al. (2021). Linkage analysis, GWAS, transcriptome analysis to identify candidate genes for rice seedlings in response to high temperature stress. BMC Plant Biol. 21, 85. doi: 10.1186/s12870-021-02857-2

PubMed Abstract | Crossref Full Text | Google Scholar

Withanawasam, D. M., Kommana, M., Pulindala, S., Eragam, A., Moode, V. N., Kolimigundla, A., et al. (2022). Improvement of Grain Yield under Moisture and Heat Stress Conditions through Marker-Assisted Pedigree Breeding in Rice (Oryza Sativa L.). Crop Pasture Sci. 73, 356–369. doi: 10.1071/CP21410

Crossref Full Text | Google Scholar

Wu, H.-C., Bulgakov, V. P., and Jinn, T.-L. (2018). Pectin methylesterases: cell wall remodeling proteins are required for plant response to heat stress. Front. Plant Sci. 9. doi: 10.3389/fpls.2018.01612

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, C., Cui, K., Wang, W., Li, Q., Fahad, S., Hu, Q., et al. (2016). Heat-induced phytohormone changes are associated with disrupted early reproductive development and reduced yield in rice. Sci. Rep. 6, 34978. doi: 10.1038/srep34978

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, C., Cui, K., Wang, W., Li, Q., Fahad, S., Hu, Q., et al. (2017). Heat-induced cytokinin transportation and degradation are associated with reduced panicle cytokinin expression and fewer spikelets per panicle in rice. Front. Plant Sci. 8. doi: 10.3389/fpls.2017.00371

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, X., Shiroto, Y., Kishitani, S., Ito, Y., and Toriyama, K. (2009). “Enhanced heat and drought tolerance in transgenic rice seedlings overexpressing osWRKY11 under the control of HSP101 promoter. Plant Cell Rep. 28, 21–305. doi: 10.1007/s00299-008-0614-x

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, Y.-S. and Yang, C.-Y. (2019). “Ethylene-mediated signaling confers thermotolerance and regulates transcript levels of heat shock factors in rice seedlings under heat stress. Botanical Stud. 60, 235. doi: 10.1186/s40529-019-0272-z

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, N., Yao, Y., Xiang, D., Du, H., Geng, Z., Yang, W., et al. (2022). A MITE variation-associated heat-inducible isoform of a heat-shock factor confers heat tolerance through regulation of JASMONATE ZIM-DOMAIN genes in rice. New Phytol. 234, 1315–1331. doi: 10.1111/nph.18068

PubMed Abstract | Crossref Full Text | Google Scholar

Wu, B., Yun, P., Zhou, H., Xia, D., Gu, Y., Li, P., et al. (2022). Natural variation in WHITE-CORE RATE 1 regulates redox homeostasis in rice endosperm to affect grain quality. Plant Cell 34, 1912–1932. doi: 10.1093/plcell/koac057

PubMed Abstract | Crossref Full Text | Google Scholar

Xalxo, R., Yadu, B., Chandra, J., Chandrakar, V., and Keshavkant, S. (2020). Alteration in carbohydrate metabolism modulates thermotolerance of plant under heat stress, in Heat stress tolerance in plants (Hoboken, New Jersey, USA: John Wiley & Sons, Ltd). doi: 10.1002/9781119432401.ch5

Crossref Full Text | Google Scholar

Xia, S., Liu, H., Cui, Y., Yu, H., Rao, Y., Yan, Y., et al. (2022). UDP-N-acetylglucosamine pyrophosphorylase enhances rice survival at high temperature. New Phytol. 233, 344–359. doi: 10.1111/nph.17768

PubMed Abstract | Crossref Full Text | Google Scholar

Xiao, J., Grandillo, S., Ahn, S. N., McCouch, S. R., Tanksley, S. D., Li, J., et al. (1996). Genes from wild rice improve yield. Nature 384, 223–224. doi: 10.1038/384223a0

Crossref Full Text | Google Scholar

Xie, Y., Le, K., Chen, X., Shen, Q., Qiu, X., Sun, L., et al. (2024). Screening of New Reference Genes for qRT-PCR Normalisation in Rice under Heat Stress. New Z. J. Crop Hortic. Sci. 52, 1–18. doi: 10.1080/01140671.2022.2072903

Crossref Full Text | Google Scholar

Xu, Y., Chu, C., and Yao, S. (2021). “The impact of high-temperature stress on rice: challenges and solutions. Crop J. 9, 963–765. doi: 10.1016/j.cj.2021.02.011

Crossref Full Text | Google Scholar

Xu, J., Henry, A., and Sreenivasulu, N. (2020). “Rice yield formation under high day and night temperatures-A prerequisite to ensure future food security. Plant Cell Environ. 43, 1595–6085. doi: 10.1111/pce.13748

PubMed Abstract | Crossref Full Text | Google Scholar

Xu, Y., Sechet, J., Wu, Y., Fu, Y., Zhu, L., Li, J., et al. (2017). Rice sucrose partitioning mediated by a putative pectin methyltransferase and homogalacturonan methylesterification1. Plant Physiol. 174, 1595–1608. doi: 10.1104/pp.16.01555

PubMed Abstract | Crossref Full Text | Google Scholar

Xu, Y., Yang, J., Wang, Y., Wang, J., Yu, Y., Long, Y., et al. (2017). OsCNGC13 promotes seed-setting rate by facilitating pollen tube growth in stylar tissues. PloS Genet. 13, e1006906. doi: 10.1371/journal.pgen.1006906

PubMed Abstract | Crossref Full Text | Google Scholar

Xu, Y., Zhang, L., Ou, S., Wang, R., Wang, Y., Chu, C., et al. (2020). Natural variations of SLG1 confer high-temperature tolerance in indica rice. Nat. Commun. 11, 5441. doi: 10.1038/s41467-020-19320-9

PubMed Abstract | Crossref Full Text | Google Scholar

Xue, W., Xing, Y., Weng, X., Zhao, Y., Tang, W., Wang, L., et al. (2008). Natural variation in ghd7 is an important regulator of heading date and yield potential in rice. Nat. Genet. 40, 761–767. doi: 10.1038/ng.143

PubMed Abstract | Crossref Full Text | Google Scholar

Yadav, M. R., Choudhary, M., Singh, J., Lal, M. K., Jha, P. K., Udawat, P., et al. (2022). Impacts, tolerance, adaptation, and mitigation of heat stress on wheat under changing climates. Int. J. Mol. Sci. 23, 5. doi: 10.3390/ijms23052838

PubMed Abstract | Crossref Full Text | Google Scholar

Yamakawa, H. and Hakata, M. (2010). “Atlas of rice grain filling-related metabolism under high temperature: joint analysis of metabolome and transcriptome demonstrated inhibition of starch accumulation and induction of amino acid accumulation. Plant Cell Physiol. 51, 795–8095. doi: 10.1093/pcp/pcq034

PubMed Abstract | Crossref Full Text | Google Scholar

Yan, C.-L., Wang, J.-B., and Li, R.-Q. (2002). Effect of heat stress on calcium ultrastructural distribution in pepper anther. Environ. Exp. Bot. 48, 161–168. doi: 10.1016/S0098-8472(02)00021-7

Crossref Full Text | Google Scholar

Yan, C., Zhan, G., Hong, X., and Yang, D. (2021). Identification and fine mapping of a major QTL, TT1-2, that plays significant roles in regulating heat tolerance in rice. Plant Mol. Biol. Rep. 39, 376–855. doi: 10.1007/s11105-020-01256-5

Crossref Full Text | Google Scholar

Yang, W., Hao, Q., Liang, J., Tan, Q., Luan, X., Lin, S., et al. (2022). Fine mapping of two major quantitative trait loci for rice chalkiness with high temperature-enhanced additive effects. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.957863

PubMed Abstract | Crossref Full Text | Google Scholar

Yang, C., Luo, A., Lu, H.-P., Davis, S. J., and Liu, J.-X. (2024). “Diurnal regulation of alternative splicing associated with thermotolerance in rice by two glycine-rich RNA-binding proteins. Sci. Bull. 69, 59–715. doi: 10.1016/j.scib.2023.11.046

PubMed Abstract | Crossref Full Text | Google Scholar

Yang, Y., Zhang, C., Zhu, D., He, H., Wei, Z., Yuan, Q., et al. (2022). Identifying candidate genes and patterns of heat-stress response in rice using a genome-wide association study and transcriptome analyses. Crop J. 10, 1633–1643. doi: 10.1016/j.cj.2022.02.011

Crossref Full Text | Google Scholar

Yao, D., Wu, J., Luo, Q., Li, J., Zhuang, W., Xiao, G., et al. (2020). Influence of high natural field temperature during grain filling stage on the morphological structure and physicochemical properties of rice (Oryza sativa L.) starch. Food Chem. 310, 125817. doi: 10.1016/j.foodchem.2019.125817

PubMed Abstract | Crossref Full Text | Google Scholar

Ye, C., Argayoso, M. A., Redoña, E. D., Sierra, S. N., Laza, M. A., Dilla, C. J., et al. (2011). Mapping QTL for heat tolerance at flowering stage in rice using SNP markers. Plant Breed. 131, 33–41. doi: 10.1111/j.1439-0523.2011.01924.x

Crossref Full Text | Google Scholar

Ye, C., Ishimaru, T., Lambio, L., Li, L., Long, Y., He, Z., et al. (2022). Marker-assisted pyramiding of QTLs for heat tolerance and escape upgrades heat resilience in rice (Oryza sativa L.). Theor. Appl. Genet. 135, 1345–1354. doi: 10.1007/s00122-022-04035-w

PubMed Abstract | Crossref Full Text | Google Scholar

Ye, C., Tenorio, F. A., Argayoso, M. A., Laza, M. A., Koh, H. J., Redoña, E. D., et al. (2015a). Identifying and confirming quantitative trait loci associated with heat tolerance at flowering stage in different rice populations. BMC Genet. 16, 41. doi: 10.1186/s12863-015-0199-7

PubMed Abstract | Crossref Full Text | Google Scholar

Ye, C., Tenorio, F. A., Redoña, E. D., Morales‑Cortezano, P. S., Cabrega, G. A., Jagadish, K. S. V., et al. (2015b). Fine-Mapping and Validating qHTSF4.1 to Increase Spikelet Fertility under Heat Stress at Flowering in Rice. TAG. Theor. Appl. Genet. Theoretische Und Angewandte Genetik 128, 1507–1517. doi: 10.1007/s00122-015-2526-9

PubMed Abstract | Crossref Full Text | Google Scholar

Ying, Y., Pang, Y., Bao, J., Ying, Y., Pang, Y., and Bao, J. (2023). Comparative ubiquitome analysis reveals diverse functions of ubiquitination in rice seed development under high-temperature stress. Seed Biol. 2. doi: 10.48130/seedbio-2023-0023

Crossref Full Text | Google Scholar

Yokotani, N., Ichikawa, T., Kondou, Y., Matsui, M., Hirochika, H., Iwabuchi, M., Oda, K., et al. (2008). Expression of rice heat stress transcription factor osHsfA2e enhances tolerance to environmental stresses in transgenic arabidopsis. Planta 227, 957–967. doi: 10.1007/s00425-007-0670-4

PubMed Abstract | Crossref Full Text | Google Scholar

Yoshida, S. (1981). Fundamentals of rice crop science. Int. Rice Res. Inst. 65–108.

Google Scholar

Zafar, S. A., Hameed, A., Ashraf, M., Khan, A. S., Qamar, Z. U., Li, X., et al. (2020). Agronomic, physiological and molecular characterisation of rice mutants revealed the key role of reactive oxygen species and catalase in high-temperature stress tolerance. Funct. Plant Biol. 47, 440–453. doi: 10.1071/FP19246

PubMed Abstract | Crossref Full Text | Google Scholar

Zaidi, S. H. R., Zakari, S. A., Zhao, Q., Khan, A. R., Shah, J. M., and Cheng, F. (2019). “Anthocyanin accumulation in black kernel mutant rice and its contribution to ROS detoxification in response to high temperature at the filling stage. Antioxidants 8, 5105. doi: 10.3390/antiox8110510

PubMed Abstract | Crossref Full Text | Google Scholar

Zang, Q., Han, X., Zhang, M., Huang, X., Jiang, M., and Huang, L. (2022). “Effects of High Temperature on Quality of Japonica Rice at Early and Middle Heading Stage under Different Planting Modes. Agronomy 12, 85. doi: 10.3390/agronomy12081833

Crossref Full Text | Google Scholar

Zhang, W., Cao, Z., Zhou, Q., Chen, J., Xu, G., Gu, J., et al. (2016). Grain filling characteristics and their relations with endogenous hormones in large- and small-grain mutants of rice. PloS One 11, e0165321. doi: 10.1371/journal.pone.0165321

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, G.-l., Chen, L.-y., Zhang, S.-t., Zheng, H., and Liu, G.-h. (2009). “Effects of high temperature stress on microscopic and ultrastructural characteristics of mesophyll cells in flag leaves of rice. Rice Sci. 16, 65–715. doi: 10.1016/S1672-6308(08)60058-X

Crossref Full Text | Google Scholar

Zhang, H., Duan, L., Dai, J. S., Zhang, C. Q., Li, J., Gu, M. H., et al. (2014). Major QTLs reduce the deleterious effects of high temperature on rice amylose content by increasing splicing efficiency of wx pre-mRNA. TAG. Theor. Appl. Genet. Theoretische Und Angewandte Genetik 127, 273–282. doi: 10.1007/s00122-013-2216-4

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, C., Li, G., Chen, T., Feng, B., Fu, W., Yan, J., et al. (2018). Heat Stress Induces Spikelet Sterility in Rice at Anthesis through Inhibition of Pollen Tube Elongation Interfering with Auxin Homeostasis in Pollinated Pistils. Rice 11, 14. doi: 10.1186/s12284-018-0206-5

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, Y., Liu, X., Su, R., Xiao, Y., Deng, H., Lu, X., et al. (2022). 9-cis-epoxycarotenoid dioxygenase 1 confers heat stress tolerance in rice seedling plants. Front. Plant Sci. 13. doi: 10.3389/fpls.2022.1092630

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, F., Liu, X., Zhang, A., Jiang, Z., Chen, L., and Zhang, X. (2019). “Genome-wide dynamic network analysis reveals a critical transition state of flower development in arabidopsis. BMC Plant Biol. 19, 115. doi: 10.1186/s12870-018-1589-6

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, Y., Tateishi‑Karimata, H., Endoh, T., Jin, Q., Li, K., Fan, X., et al. (2022). High-temperature adaptation of an osNRT2.3 allele is thermoregulated by small RNAs. Sci. Adv. 8, eadc9785. doi: 10.1126/sciadv.adc9785

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, H., Wang, X., Yan, A., Deng, J., Xie, Y., Liu, S., et al. (2023). Evolutionary analysis of respiratory burst oxidase homolog (RBOH) genes in plants and characterization of zmRBOHs. Int. J. Mol. Sci. 24, 3858. doi: 10.3390/ijms24043858

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, Q. L., Wei, Y. X., and Peng, C. L. (2018). Effects of endogenous ascorbic acid on resistance to high-temperature stress in excised rice leaves. Photosynthetica 56, 1453–1458. doi: 10.1007/s11099-018-0836-2

Crossref Full Text | Google Scholar

Zhang, B., Wu, S., Zhang, Y. E., Xu, T., Guo, F., Tang, H., et al. (2016). A high temperature-dependent mitochondrial lipase EXTRA GLUME1 promotes floral phenotypic robustness against temperature fluctuation in rice (Oryza sativa L.). PloS Genet. 12, e1006152. doi: 10.1371/journal.pgen.1006152

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, H., Xu, H., Feng, M., and Zhu, Y. (2018). “Suppression of osMADS7 in rice endosperm stabilizes amylose content under high temperature stress. Plant Biotechnol. J. 16, 18–265. doi: 10.1111/pbi.12745

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, H., Xu, H., Jiang, Y., Zhang, H., Wang, S., Wang, F., et al. (2021). Genetic control and high temperature effects on starch biosynthesis and grain quality in rice. Front. Plant Sci. 12. doi: 10.3389/fpls.2021.757997

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, S. S., Yang, H., Ding, L., Song, Z. T., Ma, H., Chang, F., et al. (2017). Tissue-specific transcriptomics reveals an important role of the unfolded protein response in maintaining fertility upon heat stress in arabidopsis. Plant Cell 29, 1007–1023. doi: 10.1105/tpc.16.00916

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, P., Yu, H., Huang, Z., Yang, P., Li, H., Huang, G., et al. (2025). Combined analysis of transcriptome and metabolome reveals the heat stress resistance of dongxiang wild rice at seedling stage. Plants 14, 1192. doi: 10.3390/plants14081192

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, X., Zhang, Q., Yang, J., Jin, Y., Wu, J., Xu, H., et al. (2023). Comparative effects of heat stress at booting and grain-filling stage on yield and grain quality of high-quality hybrid rice. Foods (Basel Switzerland) 12, 4093. doi: 10.3390/foods12224093

PubMed Abstract | Crossref Full Text | Google Scholar

Zhang, H., Zhou, J. F., Kan, Y., Shan, J. X., Ye, W. W., Dong, N. Q., et al. (2022). A genetic module at one locus in rice protects chloroplasts to enhance thermotolerance. Sci. (New York N.Y.) 376, 1293–1300. doi: 10.1126/science.abo5721

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, Q., Guan, X., Zhou, L., Asad, M. A. U. A., Xu, Y., Pan, G., et al. (2023). ABA-triggered ROS burst in rice developing anthers is critical for tapetal programmed cell death induction and heat stress-induced pollen abortion. Plant Cell Environ. 46, 1453–1471. doi: 10.1111/pce.14551

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, L., Lei, J., Huang, Y., Zhu, S., Chen, H., Huang, R., et al. (2016). Mapping quantitative trait loci for heat tolerance at anthesis in rice using chromosomal segment substitution lines. Breed. Sci. 66, 358–366. doi: 10.1270/jsbbs.15084

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, C., Liu, B., Piao, S., Wang, X., Lobell, D. B., Huang, Y., et al. (2017). Temperature increase reduces global yields of major crops in four independent estimates. Proc. Natl. Acad. Sci. 114, 9326–9331. doi: 10.1073/pnas.1701762114

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, J., Yu, X., Zhang, C., Hou, L., Wu, N., Zhang, W., et al. (2024). Harnessing microbial interactions with rice: strategies for abiotic stress alleviation in the face of environmental challenges and climate change. Sci. Total Environ. 912, 168847. doi: 10.1016/j.scitotenv.2023.168847

PubMed Abstract | Crossref Full Text | Google Scholar

Zhao, L., Zhao, C. F., Zhou, L. H., Lin, J., Zhao, Q. Y., Zhu, Z., et al. (2016). QTL mapping of dehiscence length at the basal part of thecae related to heat tolerance of rice (Oryza sativa L.). Euphytica 209, 715–723. doi: 10.1007/s10681-016-1664-8

Crossref Full Text | Google Scholar

Zheng, K., Zhao, J., Lin, D., Chen, J., Xu, J., Zhou, H., et al. (2016). The rice TCM5 gene encoding a novel deg protease protein is essential for chloroplast development under high temperatures. Rice 9, 13. doi: 10.1186/s12284-016-0086-5

PubMed Abstract | Crossref Full Text | Google Scholar

Zhou, H., Wang, X., Huo, C., Wang, H., An, Z., Sun, D., et al. (2019). A quantitative proteomics study of early heat-regulated proteins by two-dimensional difference gel electrophoresis identified osUBP21 as a negative regulator of heat stress responses in rice. Proteomics 19, e1900153. doi: 10.1002/pmic.201900153

PubMed Abstract | Crossref Full Text | Google Scholar

Zhou, H., Wang, Y., Zhang, Y., Xiao, Y., Liu, X., Deng, H., et al. (2022). Comparative analysis of heat-tolerant and heat-susceptible rice highlights the role of osNCED1 gene in heat stress tolerance. Plants 11, 8. doi: 10.3390/plants11081062

PubMed Abstract | Crossref Full Text | Google Scholar

Zhou, S., Zhu, S., Cui, S., Hou, H., Wu, H., Hao, B., et al. (2021). Transcriptional and post-transcriptional regulation of heading date in rice. New Phytol. 230, 943–956. doi: 10.1111/nph.17158

PubMed Abstract | Crossref Full Text | Google Scholar

Zhu, S., Huang, R., Wai, H. P., Xiong, H., Shen, X., He, H., et al. (2017). Mapping quantitative trait loci for heat tolerance at the booting stage using chromosomal segment substitution lines in rice. Physiol. Mol. Biol. Plants 23, 817–825. doi: 10.1007/s12298-017-0465-4

PubMed Abstract | Crossref Full Text | Google Scholar