Abscisic acid mediates the trade-off between growth and abiotic stress defense

Abstract

Global climate change impacts the yield and quality of rice, thereby restricting the sustainable development of agriculture. When plants cope with environmental stress, the interaction among signal generation, perception, and transmission and defense signal networks jointly enhances their stress resistance. Abscisic acid (ABA), as a key plant hormone, plays an important role in coordinating plant growth and adaptation to environmental stress. This review discusses the regulatory mechanisms of ABA in the growth, development, and stress response of rice; analyses the related signaling pathways, gene expression regulation, and functional characteristics under different environments; and explores how ABA balances plant growth and stress response. The research reveals the balance mechanism of ABA in rice, providing a theoretical basis for the improvement of rice varieties and the formulation of efficient cultivation strategies.

Keywords

Abiotic stress / Abscisic acid / Epigenetic regulation / Growth and development / Hormonal interactions / Rice

Abbreviations

AAO3	abscisic aldehyde oxidase 3
ABA	abscisic acid
ABC	ATP-binding cassette
ABRE	ABA-responsive element
ABT	ABA signaling termination factor
ACC	ACC synthase
APX	ascorbate peroxidase
CBF	c-repeat binding factor
CYP707A	cytochrome P450 707A
DPA	dihydrophaseic acid
DPAG	glucose conjugate
FAO	food and agriculture organization
GA	gibberellin
GE	glucose ester
GR	glutathione reductase
HSF	heat shock factors
HSP	heat-shock proteins
LEA	late embryogenesis abundant
LLPS	liquid-liquid phase separation
MITE	miniature inverted-repeat transposable element
NCED	9-cis-epoxycarotenoid dioxygenase
PA	phaseic acid
PP2C	protein phosphatase 2C
PYL	pyrabactin resistance-like
ROS	reactive oxygen species
SDGs	sustainable development goals
SLAC1	slow anion channel 1
SnRK2	SNF1-related protein kinase 2
SOS1	salt overly sensitive 1
TE	tiller enhancer
TOR	target of rapamycin
ZEP	zeaxanthin epoxidase

Authors' contribution

J.Y.Z., Z.T., C.Z., T.S., J.H.Z., and N.Y.: Writing, reviewing, and editing; J.Y.Z., Z.T., C.Z., J.L., and N.Y.: Writing original draft; J.Y.Z., T. S., and N.Y.: Conceptualization; J.Y.Z. and J.L.: Data curation; J.Y.Z.: Validation; T.S., J.H.Z., and N.Y.: Funding acquisition; J.H.Z. and N.Y.: Supervision and project administration; and N.Y.: Resources.

Availability of data and materials

Not applicable.

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Authors Jianhua Zhang and Nenghui Ye (Editorial Board members) were not involved in the journal's review nor decisions related to this manuscript.

Acknowledgements

This study was supported by the National Natural Science Foundation of China (32572250, 32171927, 32472060), Natural Science Foundation of Hunan Province (2025JJ30010, 2023NK2002), Yuelushan Laboratory Talent Program (2024RC2030), Guangdong Basic and Applied Basic Research Foundation (2022A1515111230), Shenzhen Science and Technology Program (JCYJ20220531103803008), and the Hong Kong Research Grant Council (AoE/M-403/16, AoE/M-05/12, GRF12102423, 12101722, 12105824).

References

[1] Alam I., Manghwar H., Zhang H., Yu Q., Ge L., 2022. Identification of GOLDEN2-like transcription factor genes in soybeans and their role in regulating plant development and metal ion stresses. Front. Plant Sci. 13, 1052659.

[2] Banerjee A., Roychoudhury A., 2019. Melatonin application reduces fluoride uptake and toxicity in rice seedlings by altering abscisic acid, gibberellin, auxin and antioxidant homeostasis. Plant Physiol. Biochem. 145, 164-173.

[3] Bhandari U., Gajurel A., Khadka B., Thapa I., Chand I., Bhatta D., Poudel A., Pandey M., Shrestha S., Shrestha J., 2023. Morpho-physiological and biochemical response of rice (Oryza sativa L.) to drought stress: A review. Heliyon 9, e13744.

[4] Burla B., Pfrunder S., Nagy R., Francisco R.M., Lee Y., Martinoia E., 2013. Vacuolar transport of abscisic acid glucosyl ester is mediated by ATP-binding cassette and proton-antiport mechanisms in Arabidopsis. Plant Physiol. 163, 1446-1458.

[5] Chauhan H., Khurana N., Agarwal P., Khurana P., 2011. Heat shock factors in rice (Oryza sativa L.): genome-wide expression analysis during reproductive development and abiotic stress. Mol. Genet. Genomics 286, 171-187.

[6] Chen L., Zhao Y., Xu S., Zhang Z., Xu Y., Zhang J., Chong K., 2018. OsMADS 57 together with OsTB1 coordinates transcription of its target OsWRKY94 and D14 to switch its organogenesis to defense for cold adaptation in rice New Phytol. 218, 219-231.

[7] Deng Y., Kashtoh H., Wang Q., Zhen G., Li Q., Tang L., Gao H., Zhang C., Qin L., Su M., Li F., Huang X., Wang Y., Xie Q., Clarke O.B., Hendrickson W.A., Chen Y., 2021. Structure and activity of SLAC 1 channels for stomatal signaling in leaves. Proc. Natl. Acad. Sci. U. S. A. 118, e2015151118.

[8] de Vleesschauwer D., Filipe O., Hoffman G., Seifi H.S., Haeck A., Canlas P., van Bockhaven J., de Waele E., Demeestere K., Ronald P., Hofte M., 2018. Target of rapamycin signaling orchestrates growth-defense trade-offs in plants. New Phytol. 217, 305-319.

[9] Ding Y., Yang S., 2022. Surviving and thriving: How plants perceive and respond to temperature stress. Dev. Cell 57, 947-958.

[10] Dong T., Park Y., Hwang I., 2015. Abscisic acid: biosynthesis, inactivation, homeostasis and signalling. Essays Biochem. 58, 29-48.

[11] Dong Z., Yu Y., Li S., Wang J., Tang S., Huang R., 2016. Abscisic acid antagonizes ethylene production through the ABI4-mediated transcriptional repression of ACS4 and ACS8 in Arabidopsis. Mol. Plant 9, 126-135.

[12] Dupeux F., Antoni R., Betz K., Santiago J., Gonzalez-Guzman M., Rodriguez L., Rubio S., Park S.Y., Cutler S.R., Rodriguez P.L., Ma�rquez J.A., 2011. Modulation of abscisic acid signaling in vivo by an engineered receptor-insensitive protein phosphatase type 2C allele. Plant Physiol. 156, 106-116.

[13] Estrada-Melo A.C., Ma C., Reid M.S., Jiang C., 2015. Overexpression of an ABA biosynthesis gene using a stress-inducible promoter enhances drought resistance in petunia. Hortic. Res. 2, 15013.

[14] Fatma M., Iqbal N., Gautam H., Sehar Z., Sofo A., D’Ippolito I., Khan N.A., 2021. Ethylene and sulfur coordinately modulate the antioxidant system and ABA accumulation in mustard plants under salt stress. Plants 10, 180.

[15] Feng Z., Lu G., Sun M., Jin Y., Xu Y., Liu X., Wang M., Liu M., Yang H., Guan Y., Yu T., Hu J., Xie Z., Li W., Liang Z., 2023. Comparative study of the priming effect of abscisic acid on tolerance to saline and alkaline stresses in rice seedlings. Agronomy 13, 2698.

[16] Finkelstein R.R., Lynch T.J., 2000. The Arabidopsis abscisic acid response gene ABI5 encodes a basic leucine zipper transcription factor. Plant Cell 12, 599-609.

[17] Fu D., Wu W., Mustafa G., Yang Y., Yang P., 2025. Molecular mechanisms of rice seed germination. New Crops 2, 100051.

[18] Gao W., Li M., Yang S., Gao C., Su Y., Zeng X., Jiao Z., Xu W., Zhang M., Xia K., 2022. miR 2105 and the kinase OsSAPK10 co-regulate OsbZIP86 to mediate drought-induced ABA biosynthesis in rice. Plant Physiol 189, 889-905.

[19] Geiger D., Scherzer S., Mumm P., Stange A., Marten I., Bauer H., Ache P., Matschi S., Liese A., Al-Rasheid K.A.S., Romeis T., Hedrich R., 2009. Activity of guard cell anion channel SLAC 1 is controlled by drought-stress signaling kinase-phosphatase pair. Proc. Natl. Acad. Sci. U. S. A. 106, 21425-21430.

[20] Golldack D., Li C., Mohan H., Probst N., 2014. Tolerance to drought and salt stress in plants: unraveling the signaling networks. Front. Plant Sci. 5, 151.

[21] Guo R., Wen X., Zhang W., Huang L., Peng Y., Jin L., Han H., Zhang L., Li W., Guo H., 2023a. Arabidopsis EIN2 represses ABA responses during germination and early seedling growth by inactivating HLS 1 protein independently of the canonical ethylene pathway. Plant J. 115, 1514-1527.

[22] Guo Y., Tan Y., Qu M., Hong K., Zeng L., Wang L., Zhuang C., Qian Q., Hu J., Xiong G., 2023b. OsWR2 recruits HDA 704 to regulate the deacetylation of H4K8ac in the promoter of OsABI5 in response to drought stress J. Integr. Plant Biol. 65, 1651-1669.

[23] Habibpourmehraban F., Wu Y., Masoomi-Aladizgeh F., Amirkhani A., Atwell B.J., Haynes P.A., 2023. Pre-treatment of rice plants with ABA makes them more tolerant to multiple abiotic stress. Int. J. Mol. Sci. 24, 9628.

[24] Hewage K.A.H., Yang J., Wang D., Hao G., Yang G., Zhu J., 2020. Chemical manipulation of abscisic acid signaling: a new approach to abiotic and biotic stress management in agriculture. Adv. Sci. 7, 2001265.

[25] Huang G., Kilic A., Karady M., Zhang J., Mehra P., Song X., Sturrock C.J., Zhu W., Qin H., Hartman S., Schneider H.M., Bhosale R., Dodd I.C., Sharp R.E., Huang R., Mooney S.J., Liang W., Bennett M.J., Zhang D., Pandey B.K., 2022. Ethylene inhibits rice root elongation in compacted soil via ABA and auxin-mediated mechanisms. Proc. Natl. Acad. Sci. U. S. A. 119, e2201072119.

[26] Hu Y., Han X., Yang M., Zhang M., Pan J., Yu D., 2019. The transcription factor INDUCER OF CBF EXPRESSION1 interacts with ABSCISIC ACID INSENSITIVE5 and DELLA proteins to fine-tune abscisic acid signaling during seed germination in Arabidopsis. Plant Cell 31, 1520-1538.

[27] Huizinga D.H., Omosegbon O., Omery B., Crowell D.N., 2008. Isoprenylcysteine methylation and demethylation regulate abscisic acid signaling in Arabidopsis. Plant Cell 20, 2714-2728.

[28] Imes D., Mumm P., Böhm J., Al-Rasheid K.A.S., Marten I., Geiger D., Hedrich R., 2013. Open stomata 1 (OST1) kinase controls R-type anion channel QUAC1 in Arabidopsis guard cells. Plant J. 74, 372-382.

[29] Jia M., Meng X., Song X., Zhang D., Kou L., Zhang J., Jing Y., Liu G., Liu H., Huang X., Wang Y., Yu H., Li J., 2022. Chilling-induced phosphorylation of IPA 1 by OsSAPK6 activates chilling tolerance responses in rice. Cell Discov. 8, 71.

[30] Jian L., Kang K., Choi Y., Suh M.C., Paek N.C., 2022. Mutation of OsMYB60reduces rice resilience to drought stress by attenuating cuticular wax biosynthesis. Plant J. 112, 339-351.

[31] Jiang C., Shimono M., Sugano S., Kojima M., Yazawa K., Yoshida R., Inoue H., Hayashi N., Sakakibara H., Takatsuji H., 2010. Abscisic acid interacts antagonistically with salicylic acid signaling pathway in rice-Magnaporthe grisea interaction. Mol. Plant-Microbe Interact. 23, 791-798.

[32] Jiang D., Zhou L., Chen W., Ye N., Xia J., Zhuang C., 2019. Overexpression of a microRNA-targeted NAC transcription factor improves drought and salt tolerance in Rice via ABA-mediated pathways. Rice 12, 76.

[33] Kang J., Hwang J.U., Lee M., Kim Y.Y., Assmann S.M., Martinoia E., Lee Y., 2010. PDR-type ABC transporter mediates cellular uptake of the phytohormone abscisic acid. Proc. Natl. Acad. Sci. U. S. A. 107, 2355-2360.

[34] Kavi Kishor P.B., Tiozon R.N., Fernie A.R., Sreenivasulu N., 2022. Abscisic acid and its role in the modulation of plant growth, development, and yield stability. Trends Plant Sci. 27, 1283-1295.

[35] Kumar A.R.N., Vijayalakshmi C., Vijayalakshmi D., 2015. Osmolyte accumulation, membrane stability and ABA profiles in rice genotypes exposed to heat and drought stress. Int. J. Bioresour. Stress Manage. 6, 117-122.

[36] Kuromori T., Miyaji T., Yabuuchi H., Shimizu H., Sugimoto E., Kamiya A., Moriyama Y., Shinozaki K., 2010. ABC transporter AtABCG 25 is involved in abscisic acid transport and responses. Proc. Natl. Acad. Sci. U. S. A. 107, 2361-2366.

[37] Kuromori T., Seo M., Shinozaki K., 2018. ABA transport and plant water stress responses. Trends Plant Sci. 23, 513-522.

[38] Lee S.C., Lan W., Buchanan B.B., Luan S., 2009. A protein kinase-phosphatase pair interacts with an ion channel to regulate ABA signaling in plant guard cells. Proc. Natl. Acad. Sci. U. S. A. 106, 21419-21424.

[39] Li C., Shen H., Wang T., Wang X., 2015. ABA regulates subcellular redistribution of OsABI-LIKE2, a negative regulator in ABA signaling, to control root architecture and drought resistance in Oryza sativa. Plant Cell Physiol. 56, 2396-2408.

[40] Li G., Ma Y., Wang X., Cheng N., Meng D., Chen S., Wang W., Wang X., Hu X., Yan L., Wang S., 2022. CRISPR/Cas 9 gene editing of NtAITRs, a family of transcription repressor genes, leads to enhanced drought tolerance in tobacco. Int. J. Mol. Sci. 23, 15268.

[41] Liao Y., Bai Q., Xu P., Wu T., Guo D., Peng Y., Zhang H., Deng X., Chen X., Luo M., Ali A., Wang W., Wu X., 2018. Mutation in rice abscisic acid2results in cell death, enhanced disease-resistance, altered seed dormancy and development. Front. Plant Sci. 9, 405.

[42] Liao Z., Zhang Y., Yu Q., Fang W., Chen M., Li T., Liu Y., Liu Z., Chen L., Yu S., Xia H., Xue H., Yu H., Luo L., 2023. Coordination of growth and drought responses by GA-ABA signaling in rice. New Phytol. 240, 1149-1161.

[43] Liang C., Wang Y., Zhu Y., Tang J., Hu B., Liu L., Ou S., Wu H., Sun X., Chu J., Chu C., 2014. OsNAP connects abscisic acid and leaf senescence by fine-tuning abscisic acid biosynthesis and directly targeting senescence-associated genes in rice. Proc. Natl. Acad. Sci. U. S. A. 111, 10013-10018.

[44] Liang F., Ho W.Q., Crabtree G.R., 2011. Engineering the ABA plant stress pathway for regulation of induced proximity. Sci. Signal. 4, rs2.

[45] Lim C.W., Baek W., Jung J., Kim J.H., Lee S.C., 2015. Function of ABA in stomatal defense against biotic and drought stresses. Int. J. Mol. Sci. 16, 15251-15270.

[46] Lim C.W., Lee S.C., 2020. ABA-dependent and ABA-independent functions of RCAR5/ PYL 11 in response to cold stress. Front. Plant Sci. 11, 587620.

[47] Lin Q., Wu F., Sheng P., Zhang Z., Zhang X., Guo X., Wang J., Cheng Z., Wang J., Wang H., Wan J., 2015. The SnRK2-APC/C^TE regulatory module mediates the antagonistic action of gibberellic acid and abscisic acid pathways. Nat. Commun. 6, 7981.

[48] Lin Z., Li Y., Wang Y., Liu X., Ma L., Zhang Z., Mu C., Zhang Y., Peng L., Xie S., Song C., Shi H., Zhu J., Wang P., 2021. Initiation and amplification of SnRK 2 activation in abscisic acid signaling. Nat. Commun. 12, 2456.

[49] Liu H., Si X., Wang Z., Cao L., Gao L., Zhou X., Wang W., Wang K., Jiao C., Zhuang L., Liu Y., Hou J., Li T., Hao C., Guo W., Liu J., Zhang X., 2023. TaTPP-7A positively feedback regulates grain filling and wheat grain yield through T6P-SnRK 1 signalling pathway and sugar-ABA interaction. Plant Biotechnol. J. 21, 1159-1175.

[50] Liu X., Ji P., Yang H., Jiang C., Liang Z., Chen Q., Lu F., Chen X., Yang Y., Zhang X., 2022. Priming effect of exogenous ABA on heat stress tolerance in rice seedlings is associated with the upregulation of antioxidative defense capability and heat shock-related genes. Plant Growth Regul. 98, 23-38.

[51] Lou D., Wang H., Liang G., Yu D., 2017. OsSAPK 2 confers abscisic acid sensitivity and tolerance to drought stress in rice. Front. Plant Sci. 8, 993.

[52] Lowe N.M., 2021. The global challenge of hidden hunger: perspectives from the field. Proc. Nutr. Soc. 80, 283-289.

[53] Luo X., Dai Y., Zheng C., Yang Y., Chen W., Wang Q., Chandrasekaran U., Du J., Liu W., Shu K., 2021. The ABI4-RbohD/VTC 2 regulatory module promotes reactive oxygen species (ROS) accumulation to decrease seed germination under salinity stress. New Phytol. 229, 950-962.

[54] Ma B., Yin C., He S., Lu X., Zhang W., Lu T., Chen S., Zhang J., 2014. Ethylene-induced inhibition of root growth requires abscisic acid function in rice (Oryza sativa L.) seedlings. PLoS Genet. 10, e1004701.

[55] Miao J., Li X., Li X., Tan W., You A., Wu S., Tao Y., Chen C., Wang J., Zhang D., Gong Z., Yi C., Yang Z., Gu M., Liang G., Zhou Y., 2020. OsPP2C09, a negative regulatory factor in abscisic acid signalling, plays an essential role in balancing plant growth and drought tolerance in rice. New Phytol. 227, 1417-1433.

[56] Miao R., Yuan W., Wang Y., Garcia-Maquilon I., Dang X., Li Y., Zhang J., Zhu Y., Rodriguez P.L., Xu W., 2021. Low ABA concentration promotes root growth and hydrotropism through relief of ABA INSENSITIVE 1-mediated inhibition of plasma membrane H⁺-ATPase 2. Sci. Adv. 7, eabd4113.

[57] Müller M., 2021. Foes or friends: ABA and ethylene interaction under abiotic stress. Plants 10, 448.

[58] Nakashima K., Yamaguchi-Shinozaki K., 2013. ABA signaling in stress-response and seed development. Plant Cell Reports 32, 959-970.

[59] Nakashima K., Yamaguchi-Shinozaki K., Shinozaki K., 2014. The transcriptional regulatory network in the drought response and its crosstalk in abiotic stress responses including drought, cold, and heat. Front. Plant Sci. 5, 170.

[60] Park S.Y., Fung P., Nishimura N., Jensen D.R., Fujii H., Zhao Y., Lumba S., Santiago J., Rodrigues A., Chow T.F., Alfred S.E., Bonetta D., Finkelstein R., Provart N.J., Desveaux D., Rodriguez P.L., McCourt P., Zhu J., Schroeder J.I., Volkman B.F., Cutler S.R., 2009. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science 324, 1068-1071.

[61] Park Y., Xu Z., Kim S.Y., Lee J., Choi B., Lee J., Kim H., Sim H.J., Hwang I., 2016. Spatial regulation of ABCG25, an ABA exporter, is an important component of the mechanism controlling cellular ABA levels. Plant Cell 28, 2528-2544.

[62] Qin H., Pandey B.K., Li Y., Huang G., Wang J., Quan R., Zhou J., Zhou Y., Miao Y., Zhang D., Bennett M.J., Huang R., 2022. Orchestration of ethylene and gibberellin signals determines primary root elongation in rice. Plant Cell 34, 1273-1288.

[63] Qin H., Wang J., Zhou J., Qiao J., Li Y., Quan R., Huang R., 2023. Abscisic acid promotes auxin biosynthesis to inhibit primary root elongation in rice. Plant Physiol. 191, 1953-1967.

[64] Qin P., Zhang G., Hu B., Wu J., Chen W., Ren Z., Liu Y., Xie J., Yuan H., Tu B., Ma B., Wang Y., Ye L., Li L., Xiang C., Li S., 2021. Leaf-derived ABA regulates rice seed development via a transporter-mediated and temperature-sensitive mechanism. Sci. Adv. 7, eabc8873.

[65] Qiu Q., Guo Y., Dietrich M.A., Schumaker K.S., Zhu J., 2002. Regulation of SOS1, a plasma membrane Na⁺/H⁺ exchanger in Arabidopsis thaliana, by SOS2 and SOS3. Proc. Natl. Acad. Sci. U. S. A. 99, 8436-8441.

[66] Qin Z., Lyu J., Teng Z., Meng S., Peng Y., Yuan D., Duan M., Zhang J., Ye N., 2025. ABA biosynthesis rather than ABA catabolism is induced by low temperature and inhibits seed germination by activating OsTPP3. Crop J. 13, 752-763.

[67] Rodrigues O., Reshetnyak G., Grondin A., Saijo Y., Leonhardt N., Maurel C., Verdoucq L., 2017. Aquaporins facilitate hydrogen peroxide entry into guard cells to mediate ABA and pathogen-triggered stomatal closure. Proc. Natl. Acad. Sci. U. S. A. 114, 9200-9205.

[68] Róz_ ańska E., Krępski T., Wi śniewska A., 2023. Mutations in selected ABA-related genes reduce level of Arabidopsis thaliana susceptibility to the beet cyst nematode Heterodera schachtii. Plants 12, 2299.

[69] Sah S.K., Reddy K.R., Li J., 2016. Abscisic acid and abiotic stress tolerance in crop plants. Front. Plant Sci. 7, 571.

[70] Saito S., Hirai N., Matsumoto C., Ohigashi H., Ohta D., Sakata K., Mizutani M., 2004. Arabidopsis CYP707As encode (+)-abscisic acid 8’-hydroxylase, a key enzyme in the oxidative catabolism of abscisic acid. Plant Physiol. 134, 1439-1449.

[71] Sarma B., Kashtoh H., Lama Tamang T., Bhattacharyya P.N., Mohanta Y.K., Baek K. H., 2023. Abiotic stress in rice: visiting the physiological response and its tolerance mechanisms. Plants 12, 3948.

[72] Sharp R.E., 2002. Interaction with ethylene: changing views on the role of abscisic acid in root and shoot growth responses to water stress. Plant Cell Environ. 25, 211-222.

[73] Sharp R.E., Poroyko V., Hejlek L.G., Spollen W.G., Springer G.K., Bohnert H.J., Nguyen H.T., 2004. Root growth maintenance during water deficits: physiology to functional genomics. J. Exp. Bot. 55, 2343-2351.

[74] Shu K., Zhang H., Wang S., Chen M., Wu Y., Tang S., Liu C., Feng Y., Cao X., Xie Q., 2013. ABI 4 regulates primary seed dormancy by regulating the biogenesis of abscisic acid and gibberellins in Arabidopsis. PLoS Genet. 9, e1003577.

[75] Shu K., Chen Q., Wu Y., Liu R., Zhang H., Wang P., Li Y., Wang S., Tang S., Liu C., Yang W., Cao X., Serino G., Xie Q., 2016. ABI 4 mediates antagonistic effects of abscisic acid and gibberellins at transcript and protein levels. Plant J. 85, 348-361.

[76] Song S., Wang G., Wu H., Fan X., Liang L., Zhao H., Li S., Hu Y., Liu H., Ayaad M., Xing Y., 2020. OsMFT2 is involved in the regulation of ABA signaling-mediated seed germination through interacting with OsbZIP23/66/72 in rice. Plant J. 103, 532-546.

[77] Soto-Burgos J., Bassham D.C., 2017. SnRK 1 activates autophagy via the TOR signaling pathway in Arabidopsis thaliana. PLoS One 12, e0182591.

[78] Spollen W.G., LeNoble M.E., Samuels T.D., Bernstein N., Sharp R.E., 2000. Abscisic acid accumulation maintains maize primary root elongation at low water potentials by restricting ethylene production. Plant Physiol. 122, 967-976.

[79] Sripinyowanich S., Klomsakul P., Boonburapong B., Bangyeekhun T., Asami T., Gu H., Buaboocha T., Chadchawan S., 2013. Exogenous ABA induces salt tolerance in indica rice (Oryza sativa L.): The role of OsP5CS1and OsP5CR gene expression during salt stress. Environ. Exp. Bot. 86, 94-105.

[80] Tang L., Wu A., Li S., Tuerdimaimaiti M., Zhang G., 2023. Impacts of climate change on rice grain: a literature review on what is happening, and how should we proceed? Foods 12, 536.

[81] Teng Z., Yu H., Wang G., Meng S., Liu B., Yi Y., Chen Y., Zheng Q., Liu L., Yang J., Duan M., Zhang J., Ye N., 2022b. Synergistic interaction between ABA and IAA due to moderate soil drying promotes grain filling of inferior spikelets in rice. Plant J. 109, 1457-1472.

[82] Teng Z., Chen Y., Yuan Y., Peng Y., Yi Y., Yu H., Yi Z., Yang J., Peng Y., Duan M., Zhang J., Ye N., 2022a. Identification of microRNAs regulating grain filling of rice inferior spikelets in response to moderate soil drying post-anthesis. Crop J. 10, 962-971.

[83] Teng Z., Lyu J., Chen Y., Zhang J., Ye N., 2023. Effects of stress-induced ABA on root architecture development: Positive and negative actions. Crop J. 11, 1072-1079.

[84] Theerawitaya C., Samphumphuang T., Tisarum R., Siangliw M., Cha-um S., Takabe T., Toojinda T., 2020. Expression level of Na⁺ homeostasis-related genes and salt-tolerant abilities in backcross introgression lines of rice crop under salt stress at reproductive stage. Protoplasma 257, 1595-1606.

[85] Tian X., Wang Z., Li X., Lv T., Liu H., Wang L., Niu H., Bu Q., 2015. Characterization and functional analysis of pyrabactin resistance-like abscisic acid receptor family in rice. Rice 8, 28.

[86] Tiwari S., Nutan K.K., Deshmukh R., Sarsu F., Gupta K.J., Singh A.K., Singla-Pareek S.L., Pareek A., 2022. Seedling-stage salinity tolerance in rice: Decoding the role of transcription factors. Physiol. Plant. 174, e13685.

[87] van Dijk M., Morley T., Rau M.L., Saghai Y., 2021. A meta-analysis of projected global food demand and population at risk of hunger for the period 2010-2050. Nat Food 2, 494-501.

[88] Vishwakarma K., Upadhyay N., Kumar N., Yadav G., Singh J., Mishra R.K., Kumar V., Verma R., Upadhyay R.G., Pandey M., Sharma S., 2017. Abscisic acid signaling and abiotic stress tolerance in plants: a review on current knowledge and future prospects. Front. Plant Sci. 8, 161.

[89] Waadt R., Seller C.A., Hsu P.K., Takahashi Y., Munemasa S., Schroeder J.I., 2022. Plant hormone regulation of abiotic stress responses. Nat. Rev. Mol. Cell Biol. 23, 680-694.

[90] Wang G., Li H., Wang K., Yang J., Duan M., Zhang J., Ye N., 2020a. Regulation of gene expression involved in the remobilization of rice straw carbon reserves results from moderate soil drying during grain filling. Plant J. 101, 604-618.

[91] Wang G., Li X., Ye N., Huang M., Feng L., Li H., Zhang J., 2021. OsTPP1regulates seed germination through the crosstalk with abscisic acid in rice. New Phytol. 230, 1925-1939.

[92] Wang H., Ye T., Guo Z., Yao Y., Tu H., Wang P., Zhang Y., Wang Y., Li X., Li B., Xiong H., Lai X., Xiong L., 2024. A double-stranded RNA binding protein enhances drought resistance via protein phase separation in rice. Nat. Commun. 15, 2514.

[93] Wang J., Nan N., Li N., Liu Y., Wang T., Hwang I., Liu B., Xu Z., 2020b. A DNA methylation reader-chaperone regulator-transcription factor complex activates OsHKT1;5 expression during salinity stress. Plant Cell 32, 3535-3558.

[94] Wang P., Zhao Y., Li Z., Hsu C., Liu X., Fu L., Hou Y., Du Y., Xie S., Zhang C., Gao J., Cao M., Huang X., Zhu Y., Tang K., Wang X., Tao W.A., Xiong Y., Zhu J., 2018. Reciprocal regulation of the TOR kinase and ABA receptor balances plant growth and stress response. Mol. Cell 69, 100-112.

[95] Wang T., Li X., Liu X., Yang X., Li Y., Hou B., 2023. 23 transcription factor. Plant Cell Reports 42, 17-28.

[96] Wang Z., Ren Z., Cheng C., Wang T., Ji H., Zhao Y., Deng Z., Zhi L., Lu J., Wu X., Xu S., 2020c. Counteraction of ABA-mediated inhibition of seed germination and seedling establishment by ABA signaling terminator in Arabidopsis. Mol. Plant 13, 1284-1297.

[97] Xian B., Rehmani M.S., Fan Y., Luo X., Zhang R., Xu J., Wei S., Wang L., He J., Fu A., Shu K., 2024. The ABI4-RGL 2 module serves as a double agent to mediate the antagonistic crosstalk between ABA and GA signals. New Phytol. 241, 2464-2479.

[98] Xiang H., Wang T., Zheng D., Wang L., Luo Y., Li W., 2016. Effect of ABA on seed-setting rate and physiological characteristics of rice leaves under low temperature stress at booting stage. Chin. Agric. Sci. Bull. 32, 16-23 (in Chinese with English abstract).

[99] Xie Z., Jin L., Sun Y., Zhan C., Tang S., Qin T., Liu N., 2024. OsNAC 120 balances plant growth and drought tolerance by integrating GA and ABA signaling in rice. Plant Commun. 5, 100782.

[100] Xiong H., Lu D., Li Z., Wu J., Ning X., Lin W., Bai Z., Zheng C., Sun Y., Chi W., Zhang L., Xu X., 2023. The DELLA-ABI4-HY 5 module integrates light and gibberellin signals to regulate hypocotyl elongation. Plant Commun. 4, 100597.

[101] Xu C., Shan J., Liu T., Wang Q., Ji Y., Zhang Y., Wang M., Xia N., Zhao L., 2023. CONSTANS-LIKE 1a positively regulates salt and drought tolerance in soybean. Plant Physiol. 191, 2427-2446.

[102] Xu J., Lu X., Liu Y., Lan W., Wei Z., Yu W., Li C., 2024. Interaction between ABA and NO in plants under abiotic stresses and its regulatory mechanisms. Front. Plant Sci. 15, 1330948.

[103] Xu W., Tang W., Wang C., Ge L., Sun J., Qi X., He Z., Zhou Y., Chen J., Xu Z., Ma Y., Chen M., 2020. SiMYB56confers drought stress tolerance in transgenic rice by regulating lignin biosynthesis and ABA signaling pathway. Front. Plant Sci. 11, 785.

[104] Xu Z., Kim D.H., Hwang I., 2013. ABA homeostasis and signaling involving multiple subcellular compartments and multiple receptors. Plant Cell Reports 32, 807-813.

[105] Yan L., Zhang H., Zheng Y., Cong Y., Liu C., Fan F., Zheng C., Yuan G., Pan G., Yuan D., Duan M., 2021. Transcription factor OsMADS 25 improves rice tolerance to cold stress. Hereditas 43, 1078-1087 (in Chinese with English abstract).

[106] Yang H., Li Y., Qiao Y., Sun H., Liu W., Qiao W., Li W., Liu M., Dong B., 2023. Low light stress promotes new tiller regeneration by changing source-sink relationship and activating expression of expansin genes in wheat. Plant Cell Environ. 46, 1562-1581.

[107] Yang J., Zhang J., 2018. Approach and mechanism in enhancing the remobilization of assimilates and grain-filling in rice and wheat. Chin. Sci. Bull. 63, 2932-2943 (in Chinese with English abstract).

[108] Ye N., Wang Y., Yu H., Qin Z., Zhang J., Duan M., Liu L., 2023. Abscisic acid enhances trehalose content via OsTPP3to improve salt tolerance in rice seedlings. Plants 12, 2665.

[109] You J., Chan Z., 2015. ROS regulation during abiotic stress responses in crop plants. Front. Plant Sci. 6, 1092.

[110] Yu H., Teng Z., Liu B., Lv J., Chen Y., Qin Z., Peng Y., Meng S., He Y., Duan M., Zhang J., Ye N., 2024. Transcription factor OsMYB 30 increases trehalose content to inhibit α-amylase and seed germination at low temperature. Plant Physiol. 194, 1815-1833.

[111] Zhang Q., Liu Y., Jiang Y., Li A., Cheng B., Wu J., 2022. OsASR6 enhances salt stress tolerance in rice. Int. J. Mol. Sci. 23, 9340.

[112] Zhang Y., Han L., Liu J., Chang M., Li C., Shang J., Deng Z., Tang W., Sun Y., 2025. Two E-clade protein phosphatase 2Cs enhance ABA signaling by dephosphorylating ABI 1 in Arabidopsis. Mol. Plant 18, 783-796.

[113] Zhao H., Nie K., Zhou H., Yan X., Zhan Q., Zheng Y., Song C., 2020. ABI 5 modulates seed germination via feedback regulation of the expression of the PYR/PYL/RCAR ABA receptor genes. New Phytol. 228, 596-608.

[114] Zhao J., Liu X., Wang M., Xie L., Wu Z., Yu J., Wang Y., Zhang Z., Jia Y., Liu Q., 2022. The miR528-D3module regulates plant height in rice by modulating the gibberellin and abscisic acid metabolisms. Rice 15, 27.

[115] Zhao M., Lei Y., Wu L., Qi H., Song Z., Xu M., 2024. The miR159a-PeMYB33module regulates poplar adventitious rooting through the abscisic acid signaling pathway. Plant J. 118, 879-891.

[116] Zhao X., Li C., Chen J., Ding Z., 2023. Analysis on the problems and countermeasures of cultivated land protection in southern China from the perspective of food security. Nat. Resour. Inf. 7, 30-35 (in Chinese with English abstract).

[117] Zhao Y., Xing L., Wang X., Hou Y., Gao J., Wang P., Duan C., Zhu X., Zhu J., 2014. The ABA receptor PYL 8 promotes lateral root growth by enhancing MYB77-dependent transcription of auxin-responsive genes. Sci. Signal. 7, ra53.

[118] Zhou Y., Wang Y., Li J., Liang J., 2021. In vivo FRET-FLIM reveals ER-specific increases in the ABA level upon environmental stresses. Plant Physiol. 186, 1545-1561.

[119] Zhou Z., Tang W., Sun Z., Li J., Yang B., Liu Y., Wang B., Xu D., Yang J., Zhang Y., 2023. OsCIPK 9 interacts with OsSOS3 and affects salt-related transport to improve salt tolerance. Plants 12, 3723.

[120] Zhu A., Li J., Fu W., Wang W., Tao L., Fu G., Chen T., Feng B., 2022. Abscisic acid improves rice thermo-tolerance by affecting trehalose metabolism. Int. J. Mol. Sci. 23, 10615.

[121] Zhu G., Ye N., Yang J., Peng X., Zhang J., 2011. Regulation of expression of starch synthesis genes by ethylene and ABA in relation to the development of rice inferior and superior spikelets. J. Exp. Bot. 62, 3907-3916.

[122] Zong W., Tang N., Yang J., Peng L., Ma S., Xu Y., Li G., Xiong L., 2016. Feedback regulation of ABA signaling and biosynthesis by a bZIP transcription factor targets drought-resistance-related genes. Plant Physiol. 171, 2810-2825.