[1]

Kan Y, Mu XR, Gao J, Lin HX, Lin Y. 2023. The molecular basis of heat stress responses in plants. Molecular Plant 16:1612−1634

doi: 10.1016/j.molp.2023.09.013
[2]

Matthews HD, Wynes S. 2022. Current global efforts are insufficient to limit warming to 1.5 °C. Science 376:1404−1409

doi: 10.1126/science.abo3378
[3]

Zhao C, Liu B, Piao S, Wang X, Lobell DB, et al. 2017. Temperature increase reduces global yields of major crops in four independent estimates. Proceedings of the National Academy of Sciences of the United States of America 114:9326−9331

doi: 10.1073/pnas.1701762114
[4]

Heikonen S, Heino M, Jalava M, Siebert S, Viviroli D, et al. 2025. Climate change threatens crop diversity at low latitudes. Nature Food 6:331−342

doi: 10.1038/s43016-025-01135-w
[5]

Li JY, Yang C, Xu J, Lu HP, Liu JX. 2023. The hot science in rice research: how rice plants cope with heat stress. Plant, Cell & Environment 46:1087−1103

doi: 10.1111/pce.14509
[6]

Matsui T, Kobayasi K, Yoshimoto M, Hasegawa T, Tian X. 2020. Dependence of pollination and fertilization in rice (Oryza sativa L.) on floret height within the canopy. Field Crops Research 249:107741

doi: 10.1016/j.fcr.2020.107741
[7]

Liu M, Zhou Y, Sun J, Mao F, Yao Q, et al. 2023. From the floret to the canopy: high temperature tolerance during flowering. Plant Communications 4:100629

doi: 10.1016/j.xplc.2023.100629
[8]

Liu J, Wu Y, Pan X, Su Q, Liu Z, et al. 2025. Research progress on heat stress tolerance mechanisms in rice during flowering stage. Scientia Sinica Vitae 55:2212−2222

doi: 10.1360/SSV-2025-0215
[9]

Xing YH, Lu H, Zhu X, Deng Y, Xie Y, et al. 2024. How rice responds to temperature changes and defeats heat stress. Rice 17:73

doi: 10.1186/s12284-024-00748-2
[10]

Xia M, Qi H. 2004. Effects of high-temperature damage on seed setting rate of hybrid combinations derived from four male sterile lines. Hubei Agricultural Sciences 2004:21−22
[11]

Yang H, Huang Z, Jiang Z, Wang X. 2004. Heat damage during flowering period and its prevention techniques for early and medium rice in Anhui in 2003. Journal of Anhui Agricultural Sciences 1:3−4 (in Chinese)

doi: 10.13989/j.cnki.0517-6611.2004.01.002
[12]

Wang HY. 2016. Investigation, analysis and prevention countermeasures of rice heat damage- a case study of Ma'anshan city in 2013. Journal of Anhui Agricultural Sciences 8:50−52
[13]

Luo Z, Yang Y, Tang Y, Fan L. 2011. Study on characteristics of heat damage on rice in Chongqing in context of climatic change. Southwest China Journal of Agricultural Sciences 6:2185−2189

doi: 10.3969/j.issn.1001-4829.2011.06.033
[14]

Song Y, Wang C, Linderholm HW, Fu Y, Cai W, et al. 2022. The negative impact of increasing temperatures on rice yields in southern China. Science of the Total Environment 820:153262

doi: 10.1016/j.scitotenv.2022.153262
[15]

Wang Y, Wang L, Zhou J, Hu S, Chen H, et al. 2019. Research progress on heat stress of rice at flowering stage. Rice Science 26:1−10

doi: 10.1016/j.rsci.2018.06.009
[16]

Shen C, Zhang Y, Li G, Shi J, Wang D, et al. 2023. MADS8 is indispensable for female reproductive development at high ambient temperatures in cereal crops. The Plant Cell 36:65−84

doi: 10.1093/plcell/koad246
[17]

Zhang B, Wu S, Zhang YE, Xu T, Guo F, et al. 2016. A high temperature-dependent mitochondrial lipase EXTRA GLUME1 promotes floral phenotypic robustness against temperature fluctuation in rice (Oryza sativa L.). PLoS Genetics 12:e1006152

doi: 10.1371/journal.pgen.1006152
[18]

Zhang P, Zhu W, He Y, Fan J, Shi J, et al. 2023. THERMOSENSITIVE BARREN PANICLE (TAP) is required for rice panicle and spikelet development at high ambient temperature. New Phytologist 237:855−869

doi: 10.1111/nph.18551
[19]

Cai Z, Wang G, Li J, Kong L, Tang W, et al. 2023. Thermo-Sensitive Spikelet Defects 1 acclimatizes rice spikelet initiation and development to high temperature. Plant Physiology 191:1684−1701

doi: 10.1093/plphys/kiac576
[20]

Dong K, Wu F, Cheng S, Li S, Zhang F, et al. 2024. OsPRMT6a-mediated arginine methylation of OsJAZ1 regulates jasmonate signaling and spikelet development in rice. Molecular Plant 17:900−919

doi: 10.1016/j.molp.2024.04.014
[21]

Song X, Wang D, Ma L, Chen Z, Li P, et al. 2012. Rice RNA-dependent RNA polymerase 6 acts in small RNA biogenesis and spikelet development. The Plant Journal 71:378−389

doi: 10.1111/j.1365-313X.2012.05001.x
[22]

Si F, Yang C, Yan B, Yan W, Tang S, et al. 2022. Control of OsARF3a by OsKANADI1 contributes to lemma development in rice. The Plant Journal 110:1717−1730

doi: 10.1111/tpj.15766
[23]

Gu X, Si F, Feng Z, Li S, Liang D, et al. 2023. The OsSGS3-tasiRNA-OsARF3 module orchestrates abiotic-biotic stress response trade-off in rice. Nature Communications 14:4441

doi: 10.1038/s41467-023-40176-2
[24]

Yao Q, Li P, Wang X, Liao S, Wang P, et al. 2024. Molecular mechanisms underlying the negative effects of transient heatwaves on crop fertility. Plant Communications 5:101009

doi: 10.1016/j.xplc.2024.101009
[25]

Shi H, Chan Z. 2014. Improvement of plant abiotic stress tolerance through modulation of the polyamine pathway. Journal of Integrative Plant Biology 56:114−121

doi: 10.1111/jipb.12128
[26]

Zhou R, Hu Q, Pu Q, Chen M, Zhu X, et al. 2020. Spermidine enhanced free polyamine levels and expression of polyamine biosynthesis enzyme gene in rice spikelets under heat tolerance before heading. Scientific Reports 10:8976

doi: 10.1038/s41598-020-64978-2
[27]

Feng B, Zhang C, Chen T, Zhang X, Tao L, et al. 2018. Salicylic acid reverses pollen abortion of rice caused by heat stress. BMC Plant Biology 18:245

doi: 10.1186/s12870-018-1472-5
[28]

Rezaul IM, Feng B, Chen T, Fu W, Zhang C, 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
[29]

Zhou H, Wang Y, Zhang Y, Xiao Y, Liu X, 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):1062

doi: 10.3390/plants11081062
[30]

Zhao Q, Guan X, Zhou L, Asad MAU, Xu Y, 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 & Environment 46:1453−1471

doi: 10.1111/pce.14551
[31]

Zhao Q, Guan X, Zhou L, Xu Y, Asad MAU, et al. 2023. OsPDIL1-1 controls ROS generation by modulating NADPH oxidase in developing anthers to alter the susceptibility of floret fertility to heat for rice. Environmental and Experimental Botany 205:105103

doi: 10.1016/j.envexpbot.2022.105103
[32]

Cao Z, Tang H, Cai Y, Zeng B, Zhao J, 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 Biotechnology Journal 20:1591−1605

doi: 10.1111/pbi.13835
[33]

Lin S, Liu Z, Sun S, Xue F, Li H, et al. 2023. Rice HEAT SHOCK PROTEIN60-3B maintains male fertility under high temperature by starch granule biogenesis. Plant Physiology 192:2301−2317

doi: 10.1093/plphys/kiad136
[34]

Mao X, Yu H, Xue J, Zhang L, Zhu Q, et al. 2025. OsRHS negatively regulates rice heat tolerance at the flowering stage by interacting with the HSP protein cHSP70-4. Plant, Cell & Environment 48:350−364

doi: 10.1111/pce.15152
[35]

Peng G, Liu Z, Zhuang C, Zhou H. 2023. Environment-sensitive genic male sterility in rice and other plants. Plant, Cell & Environment 46:1120−1142

doi: 10.1111/pce.14503
[36]

Cai Z, Xu C, Liu X, Lv Y, Ouyang Y, et al. 2024. Exploiting male sterility toward the development of hybrid rice. Seed Biology 3:e019

doi: 10.48130/seedbio-0024-0018
[37]

Zhou H, Liu Q, Li J, Jiang D, Zhou L, et al. 2012. Photoperiod- and thermo-sensitive genic male sterility in rice are caused by a point mutation in a novel noncoding RNA that produces a small RNA. Cell Research 22:649−660

doi: 10.1038/cr.2012.28
[38]

Ding J, Lu Q, Ouyang Y, Mao H, Zhang P, et al. 2012. A long noncoding RNA regulates photoperiod-sensitive male sterility, an essential component of hybrid rice. Proceedings of the National Academy of Sciences of the United States of America 109:2654−2659

doi: 10.3410/f.14267349.15779554
[39]

Zhou H, He M, Li J, Chen L, Huang Z, et al. 2016. Development of commercial thermo-sensitive genic male sterile rice accelerates hybrid rice breeding using the CRISPR/Cas9-mediated TMS5 editing system. Scientific Reports 6:37395

doi: 10.1038/srep37395
[40]

Zhou H, Zhou M, Yang Y, Li J, Zhu L, et al. 2014. RNase ZS1 processes UbL40 mRNAs and controls thermosensitive genic male sterility in rice. Nature Communications 5:4884

doi: 10.1038/ncomms5884
[41]

Yan B, Liu C, Sun J, Mao Y, Zhou C, et al. 2024. Impaired 2', 3' -cyclic phosphate tRNA repair causes thermo-sensitive genic male sterility in rice. Cell Research 34:763−775

doi: 10.1038/s41422-024-01012-4
[42]

Wu L, Jing X, Zhang B, Chen S, Xu R, et al. 2022. A natural allele of OsMS1 responds to temperature changes and confers thermosensitive genic male sterility. Nature Communications 13:2055

doi: 10.1038/s41467-022-29648-z
[43]

Yu J, Han J, Kim YJ, Song M, Yang Z, et al. 2017. Two rice receptor-like kinases maintain male fertility under changing temperatures. Proceedings of the National Academy of Sciences of the United States of America 114:12327−12332

doi: 10.1073/pnas.1705189114
[44]

Chen R, Zhao X, Shao Z, Wei Z, Wang Y, et al. 2007. Rice UDP-glucose pyrophosphorylase1 is essential for pollen callose deposition and its cosuppression results in a new type of thermosensitive genic male sterility. The Plant Cell 19:847−861

doi: 10.1105/tpc.106.044123
[45]

Han Y, Jiang SZ, Zhong X, Chen X, Ma CK, et al. 2023. Low temperature compensates for defective tapetum initiation to restore the fertility of the novel TGMS line ostms15. Plant Biotechnology Journal 21:1659−1670

doi: 10.1111/pbi.14066
[46]

Zhang YF, Li YL, Zhong X, Wang JJ, Zhou L, et al. 2022. Mutation of glucose-methanol-choline oxidoreductase leads to thermosensitive genic male sterility in rice and Arabidopsis. Plant Biotechnology Journal 20:2023−2035

doi: 10.1111/pbi.13886
[47]

Zhou L, Mao YC, Yang YM, Wang JJ, Zhong X, et al. 2024. Temperature and light reverse the fertility of rice P/TGMS line ostms19 via reactive oxygen species homeostasis. Plant Biotechnology Journal 22:2020−2032

doi: 10.1111/pbi.14322
[48]

Peng G, Liu M, Zhu L, Luo W, Wang Q, et al. 2023. The E3 ubiquitin ligase CSIT1 regulates critical sterility-inducing temperature by ribosome-associated quality control to safeguard two-line hybrid breeding in rice. Molecular Plant 16:1695−1709

doi: 10.1016/j.molp.2023.09.016
[49]

Peng G, Liu M, Luo Z, Deng S, Wang Q, et al. 2024. An E3 ubiquitin ligase CSIT2 controls critical sterility-inducing temperature of thermo-sensitive genic male sterile rice. New Phytologist 241:2059−2074

doi: 10.1111/nph.19520
[50]

Liu W, Li J, Sun J, Liu C, Yan B, et al. 2025. The E3 ligase OsHel2 impedes readthrough of stalled mRNAs to regulate male fertility in thermosensitive genic male sterile rice. Plant Communications 6:101192

doi: 10.1016/j.xplc.2024.101192
[51]

Zhou C, Yan B, Liu C, Sun J, Yang Y, et al. 2025. The OsRqc1-OsVms1 module regulates the temperature threshold in thermo-sensitive genic male-sterile rice lines. Science China-Life Sciences 68:3074−3087

doi: 10.1007/s11427-025-3018-9
[52]

Zhou C, Liu C, Yan B, Sun J, Li S, et al. 2025. TRNA selectivity during ribosome-associated quality control regulates the critical sterility-inducing temperature in two-line hybrid rice. Proceedings of the National Academy of Sciences of the United States of America 122:e2507697122

doi: 10.1073/pnas.2507697122
[53]

Mineri L, Bono GA, Sergi E, Colleoni PE, Morandini P, et al. 2025. OsMAINTENANCE OF MERISTEM LIKE 1 controls style number at high temperatures in rice. Plant Molecular Biology 115:24

doi: 10.1007/s11103-025-01553-1
[54]

Li H, You C, Yoshikawa M, Yang X, Gu H, et al. 2022. A spontaneous thermo-sensitive female sterility mutation in rice enables fully mechanized hybrid breeding. Cell Research 32:931−945

doi: 10.1038/s41422-022-00711-0
[55]

Zhang Z, Guo YY, Wang YC, Zhou L, Fan J, et al. 2024. A point mutation in the meiotic crossover formation gene HEI10/TFS2 leads to thermosensitive genic sterility in rice. The Plant Journal 118:506−518

doi: 10.1111/tpj.16621
[56]

Krishna Jagadish SV. 2020. Heat stress during flowering in cereals – effects and adaptation strategies. New Phytologist 226:1567−1572

doi: 10.1111/nph.16429
[57]

Bheemanahalli R, Sathishraj R, Manoharan M, Sumanth HN, Muthurajan R, et al. 2017. Is early morning flowering an effective trait to minimize heat stress damage during flowering in rice? Field Crops Research 203:238−242

doi: 10.1016/j.fcr.2016.11.011
[58]

Song R, Wang M, Yao Q, Han Y, Liu P, et al. 2025. Shifting flower timing to beat the heat of warming climate in crops. Seed Biology 4:e013

doi: 10.48130/seedbio-0025-0013
[59]

Wang M, Zhu X, Peng G, Liu M, Zhang S, et al. 2022. Methylesterification of cell-wall pectin controls the diurnal flower-opening times in rice. Molecular Plant 15:956−972

doi: 10.1016/j.molp.2022.04.004
[60]

Xu P, Wu T, Ali A, Zhang H, Liao Y, et al. 2022. EARLY MORNING FLOWERING1 (EMF1) regulates the floret opening time by mediating lodicule cell wall formation in rice. Plant Biotechnology Journal 20:1441−1443

doi: 10.1111/pbi.13860
[61]

Wang M, Zhu X, Huang Z, Chen M, Xu P, et al. 2024. Controlling diurnal flower-opening time by manipulating the jasmonate pathway accelerates development of indica-japonica hybrid rice breeding. Plant Biotechnology Journal 22:2267−2281

doi: 10.1111/pbi.14343
[62]

Ding W, Gou Y, Li Y, Li J, Fang Y, et al. 2024. A jasmonate-mediated regulatory network modulates diurnal floret opening time in rice. New Phytologist 244:176−191

doi: 10.1111/nph.20039
[63]

Zhu X, Wang M, Huang Z, Chen M, Xu P, et al. 2024. The OsMYC2–JA feedback loop regulates diurnal flower-opening time via cell wall loosening in rice. The Plant Journal 119:2585−2598

doi: 10.1111/tpj.16910
[64]

Gou Y, Heng Y, Ding W, Xu C, Tan Q, et al. 2024. Natural variation in OsMYB8 confers diurnal floret opening time divergence between indica and japonica subspecies. Nature Communications 15:2262

doi: 10.1038/s41467-024-46579-z
[65]

Wang Y, Lv X, Sheng D, Hou X, Mandal S, et al. 2023. Heat-dependent postpollination limitations on maize pollen tube growth and kernel sterility. Plant, Cell & Environment 46:3822−3838

doi: 10.1111/pce.14702
[66]

Zhou L, Asad MAU, Guan X, Pan G, Zhang Y, et al. 2024. Rice myo-inositol-3-phosphate synthase 2 (RINO2) alleviates heat injury-induced impairment in pollen germination and tube growth by modulating ca2+ signaling and actin filament cytoskeleton. The Plant Journal 119:861−878

doi: 10.1111/tpj.16802
[67]

Nakata M, Fukamatsu Y, Miyashita T, Hakata M, Kimura R, et al. 2017. High temperature-induced expression of rice α-amylases in developing endosperm produces chalky grains. Frontiers in Plant Science 8:2089

doi: 10.3389/fpls.2017.02089
[68]

Takehara K, Murata K, Yamaguchi T, Yamaguchi K, Chaya G, et al. 2018. Thermo-responsive allele of sucrose synthase 3 (sus3) provides high-temperature tolerance during the ripening stage in rice (Oryza sativa L.). Breeding Science 68:336−342

doi: 10.1270/jsbbs.18007
[69]

Lou H, Li S, Shi Z, Zou Y, Zhang Y, et al. 2025. Engineering source-sink relations by prime editing confers heat-stress resilience in tomato and rice. Cell 188:530−549.e20

doi: 10.1016/j.cell.2024.11.005
[70]

Tabassum R, Dosaka T, Ichida H, Morita R, Ding Y, et al. 2020. FLOURY ENDOSPERM11-2 encodes plastid HSP70-2 involved with the temperature-dependent chalkiness of rice (Oryza sativa L.) grains. The Plant Journal 103:604−616

doi: 10.1111/tpj.14752
[71]

Lu HP, Liu XH, Wang MJ, Zhu QY, Lyu YS, et al. 2025. The NAT1–bHLH110–CER1/CER1l module regulates heat stress tolerance in rice. Nature Genetics 57:427−440

doi: 10.1038/s41588-024-02065-2
[72]

Wu H, Ren Y, Dong H, Xie C, Zhao L, et al. 2024. FLOURY ENDOSPERM24, a heat shock protein 101 (HSP101), is required for starch biosynthesis and endosperm development in rice. New Phytologist 242:2635−2651

doi: 10.1111/nph.19761
[73]

Lu F, Jiao G, Qiu J, Zhao S, Zhao F, et al. 2025. A molecular module improves rice grain quality and yield at high temperatures. National Science Review 12:nwae416

doi: 10.1093/nsr/nwae416
[74]

Li W, Yang K, Hu C, Abbas W, Zhang J, et al. 2025. A natural gene on-off system confers field thermotolerance for grain quality and yield in rice. Cell 188:3661−3678.e21

doi: 10.1016/j.cell.2025.04.011
[75]

Liu X, Zhong X, Liao J, Ji P, Yang J, et al. 2023. Exogenous abscisic acid improves grain filling capacity under heat stress by enhancing antioxidative defense capability in rice. BMC Plant Biology 23:619

doi: 10.1186/s12870-023-04638-5
[76]

Qin P, Zhang G, Hu B, Wu J, Chen W, et al. 2021. Leaf-derived ABA regulates rice seed development via a transporter-mediated and temperature-sensitive mechanism. Science Advances 7(3):eabc8873

doi: 10.1126/sciadv.abc8873
[77]

Wu B, Yun P, Zhou H, Xia D, Gu Y, et al. 2022. Natural variation in WHITE-CORE RATE 1 regulates redox homeostasis in rice endosperm to affect grain quality. The Plant Cell 34:1912−1932

doi: 10.1093/plcell/koac057
[78]

He W, Li W, Luo X, Tang Y, Wang L, et al. 2023. Rice FERONIA-LIKE RECEPTOR 3 and 14 affect grain quality by regulating redox homeostasis during endosperm development. Journal of Experimental Botany 74:3003−3018

doi: 10.1093/jxb/erad077
[79]

Sandhu J, Irvin L, Chandaran AK, Oguro S, Paul P, et al. 2024. Natural variation in LONELY GUY-like 1 regulates rice grain weight under warmer night conditions. Plant Physiology 196:164−180

doi: 10.1093/plphys/kiae313
[80]

Chen Z, Zhou W, Guo X, Ling S, Li 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
[81]

Chen C, Begcy K, Liu K, Folsom JJ, Wang Z, et al. 2016. Heat stress yields a unique MADS box transcription factor in determining seed size and thermal sensitivity. Plant Physiology 171:606−622

doi: 10.1104/pp.15.01992
[82]

El-kereamy A, Bi YM, Ranathunge K, Beatty PH, Good AG, et al. 2012. The rice R2R3-MYB transcription factor OsMYB55 is involved in the tolerance to high temperature and modulates amino acid metabolism. PLoS One 7:e52030

doi: 10.1371/journal.pone.0052030
[83]

Ren Y, Huang Z, Jiang H, Wang Z, Wu F, et al. 2021. A heat stress responsive NAC transcription factor heterodimer plays key roles in rice grain filling. Journal of Experimental Botany 72:2947−2964

doi: 10.1093/jxb/erab027
[84]

Xu H, Li X, Zhang H, Wang L, Zhu Z, et al. 2020. High temperature inhibits the accumulation of storage materials by inducing alternative splicing of OsbZIP58 during filling stage in rice. Plant, Cell & Environment 43:1879−1896

doi: 10.1111/pce.13779
[85]

Cao R, Zhao S, Jiao G, Duan Y, Ma L, et al. 2022. OPAQUE3, encoding a transmembrane bZIP transcription factor, regulates endosperm storage protein and starch biosynthesis in rice. Plant Communications 3:100463

doi: 10.1016/j.xplc.2022.100463
[86]

Yuan H, Zeng J, Xu Z, Yuan M, Zhou X, et al. 2025. OsPPR8, a pentatricopeptide repeat protein, regulates splicing of mitochondrial nad2 intron 3 to affect grain quality and high-temperature tolerance in rice. The Plant Journal 122:e70246

doi: 10.1111/tpj.70246
[87]

Folsom JJ, Begcy K, Hao X, Wang D, Walia H. 2014. Rice Fertilization-Independent Endosperm1 regulates seed size under heat stress by controlling early endosperm development. Plant Physiology 165:238−248

doi: 10.1104/pp.113.232413
[88]

Li G, Cao R, Ma L, Jiao G, Chen P, et al. 2023. OsLEA1b modulates starch biosynthesis at high temperatures in rice. Plants 12(23):4070

doi: 10.3390/plants12234070
[89]

Guo SQ, Chen YX, Ju YL, Pan CY, Shan JX, et al. 2025. Fine-tuning gibberellin improves rice alkali–thermal tolerance and yield. Nature 639:162−171

doi: 10.1038/s41586-024-08486-7
[90]

Kan Y, Mu XR, Zhang H, Gao J, Shan JX, et al. 2022. TT2 controls rice thermotolerance through SCT1-dependent alteration of wax biosynthesis. Nature Plants 8:53−67

doi: 10.1038/s41477-021-01039-0
[91]

Kan Y, Mu XR, Qu F, Dai ZA, Gao J, et al. 2025. A stepwise decoding mechanism for heat sensing in plants connects lipid remodeling to a nuclear signaling cascade. Cell 188(26):7378−7396.e23

doi: 10.1016/j.cell.2025.11.003
[92]

Zhang H, Zhou JF, Kan Y, Shan JX, Ye WW, et al. 2022. A genetic module at one locus in rice protects chloroplasts to enhance thermotolerance. Science 376:1293−1300

doi: 10.1126/science.abo5721
[93]

Lu J, Chen J, Chen H, Fan Z, Lin L, et al. 2026. Heat shock transcription factor OsHsfc1a enhances rice seedling thermotolerance by regulating OsMFT1 and preserving chloroplast structure under heat stress. Plant Biotechnology Journal 24(3):1898−1915

doi: 10.1111/pbi.70458
[94]

Ambavaram MMR, Basu S, Krishnan A, Ramegowda V, Batlang U, et al. 2014. Coordinated regulation of photosynthesis in rice increases yield and tolerance to environmental stress. Nature Communications 5:5302

doi: 10.1038/ncomms6302
[95]

Liu J, Li X, Wang K, Wang T, Meng Y, et al. 2025. The splicing auxiliary factor OsU2AF35a enhances thermotolerance via protein separation and promoting proper splicing of OsHSA32 pre-mRNA in rice. Plant Biotechnology Journal 23:1308−1328

doi: 10.1111/pbi.14587
[96]

Yang C, Luo A, Lu HP, Davis SJ, Liu JX. 2024. Diurnal regulation of alternative splicing associated with thermotolerance in rice by two glycine-rich RNA-binding proteins. Science Bulletin 69:59−71

doi: 10.1016/j.scib.2023.11.046
[97]

Qiu Z, Kang S, He L, Zhao J, Zhang S, et al. 2018. The newly identified heat-stress sensitive albino 1 gene affects chloroplast development in rice. Plant Science 267:168−179

doi: 10.1016/j.plantsci.2017.11.015
[98]

Wang D, Qin B, Li X, Tang D, Zhang YE, et al. 2016. Nucleolar DEAD-box RNA helicase TOGR1 regulates thermotolerant growth as a pre-rRNA chaperone in rice. PLoS Genetics 12:e1005844

doi: 10.1371/journal.pgen.1005844
[99]

Wang J, Xu J, Wang L, Zhou M, Nian J, et al. 2023. SEMI-ROLLED LEAF 10 stabilizes catalase isozyme B to regulate leaf morphology and thermotolerance in rice (Oryza sativa L.). Plant Biotechnology Journal 21:819−838

doi: 10.1111/pbi.13999
[100]

Chen K, Guo T, Li XM, Zhang YM, Yang YB, et al. 2019. Translational regulation of plant response to high temperature by a dual-function tRNAHis guanylyltransferase in rice. Molecular Plant 12:1123−1142

doi: 10.1016/j.molp.2019.04.012
[101]

Li XM, Chao DY, Wu Y, Huang X, Chen K, et al. 2015. Natural alleles of a proteasome α2 subunit gene contribute to thermotolerance and adaptation of African rice. Nature Genetics 47:827−833

doi: 10.1038/ng.3305
[102]

Yu HX, Cao YJ, Yang YB, Shan JX, Ye WW, et al. 2024. A TT1–SCE1 module integrates ubiquitination and SUMOylation to regulate heat tolerance in rice. Molecular Plant 17:1899−1918

doi: 10.1016/j.molp.2024.11.007
[103]

Liu X, Ji P, Liao J, Duan X, Luo Z, et al. 2025. CRISPR/Cas knockout of the NADPH oxidase gene OsRbohB reduces ROS overaccumulation and enhances heat stress tolerance in rice. Plant Biotechnology Journal 23:336−351

doi: 10.1111/pbi.14500
[104]

El-Esawi MA, Alayafi AA. 2019. Overexpression of rice Rab7 gene improves drought and heat tolerance and increases grain yield in rice (Oryza sativa L.). Genes 10(1):56

doi: 10.3390/genes10010056
[105]

Dong J, Chen K, Chen L, Zheng M, Nie S, et al. 2025. OsNRAMP7 positively regulates heat tolerance at seedling and reproductive stages in rice. Plant Stress 16:100870

doi: 10.1016/j.stress.2025.100870
[106]

Avsec Ž, Latysheva N, Cheng J, Novati G, Taylor KR, et al. 2026. Advancing regulatory variant effect prediction with AlphaGenome. Nature 649:1206−1218

doi: 10.1038/s41586-025-10014-0
[107]

Vamathevan J, Clark D, Czodrowski P, Dunham I, Ferran E, et al. 2019. Applications of machine learning in drug discovery and development. Nature Reviews Drug Discovery 18:463−477

doi: 10.1038/s41573-019-0024-5
[108]

Jumper J, Evans R, Pritzel A, Green T, Figurnov M, et al. 2021. Highly accurate protein structure prediction with AlphaFold. Nature 596:583−589

doi: 10.1038/s41586-021-03819-2
[109]

Alipanahi B, Delong A, Weirauch MT, Frey BJ. 2015. Predicting the sequence specificities of DNA- and RNA-binding proteins by deep learning. Nature Biotechnology 33:831−838

doi: 10.1038/nbt.3300