Academic Journal
Προβολή σε EDS

Exogenous abscisic acid enhances foxtail millet grain weight under drought stress: Insights from integrative transcriptomics and weighted gene co-expression network analysis.

Λεπτομέρειες βιβλιογραφικής εγγραφής
Τίτλος: Exogenous abscisic acid enhances foxtail millet grain weight under drought stress: Insights from integrative transcriptomics and weighted gene co-expression network analysis.
Συγγραφείς: Guo S; The Research Center of Soil and Water Conservation and Ecological Environment, Chinese Academy of Sciences and Ministry of Education, Yangling, China.; State Key Laboratory of Soil and Water Conservation and Desertification Control, Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resources, Yangling, China.; University of Chinese Academy of Sciences, Beijing, China., Han F; College of Resources and Environment, Shandong Agricultural University, Taian, China., Yan J; Shaanxi Key Laboratory of Ecological Restoration in Shanbei Mining Area, College of Life Science, Yulin University, Yulin, China., Li H; State Key Laboratory of Soil and Water Conservation and Desertification Control, Northwest A&F University, Yangling, China., Chen Y; School of Agriculture and Environment, The University of Western Australia, Perth, Western Australia, Australia., Yin L; State Key Laboratory of Soil and Water Conservation and Desertification Control, Northwest A&F University, Yangling, China., Zhang S; The Research Center of Soil and Water Conservation and Ecological Environment, Chinese Academy of Sciences and Ministry of Education, Yangling, China.; State Key Laboratory of Soil and Water Conservation and Desertification Control, Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resources, Yangling, China.; University of Chinese Academy of Sciences, Beijing, China.; State Key Laboratory of Soil and Water Conservation and Desertification Control, Northwest A&F University, Yangling, China.
Πηγή: Journal of the science of food and agriculture [J Sci Food Agric] 2026 Aug 30; Vol. 106 (11), pp. 6672-6685. Date of Electronic Publication: 2026 May 08.
Τύπος έκδοσης: Journal Article
Γλώσσα: English
Στοιχεία περιοδικού: Publisher: John Wiley & Sons Country of Publication: England NLM ID: 0376334 Publication Model: Print-Electronic Cited Medium: Internet ISSN: 1097-0010 (Electronic) Linking ISSN: 00225142 NLM ISO Abbreviation: J Sci Food Agric Subsets: MEDLINE
Imprint Name(s): Publication: <2005-> : Chichester, West Sussex : John Wiley & Sons
Original Publication: London, Society of Chemical Industry.
Ιατρικοί όροι (MeSH): Abscisic Acid*/pharmacology , Setaria Plant*/genetics , Setaria Plant*/growth & development , Setaria Plant*/drug effects , Setaria Plant*/metabolism , Plant Proteins*/genetics , Plant Proteins*/metabolism , Seeds*/growth & development , Seeds*/genetics , Seeds*/drug effects , Seeds*/metabolism , Plant Growth Regulators*/pharmacology, Gene Expression Regulation, Plant/drug effects ; Transcriptome/drug effects ; Droughts ; Gene Expression Profiling ; Drought Resistance ; Stress, Physiological ; Gene Regulatory Networks
Περίληψη: Background: This study aimed to explore the underlying regulatory mechanism by which exogenous abscisic acid (ABA) application influences foxtail millet grain weight under drought stress at the heading-flowering stage. A two-year field experiment was conducted with two irrigation treatments (well-watered (CK) and drought stress (DS)) and four ABA concentrations (0, 10, 20, and 30 mg L-1, designated as T1-T4). The grain filling rate and grain weight were measured, and transcriptome sequencing and weighted gene co-expression network analysis (WGCNA) were performed at three stages to select candidate genes regulating grain weight.
Results: The grain weight exhibited an S-shaped growth pattern during the grain-filling period. Compared with CK, drought stress significantly reduced the grain-filling rate and grain weight, decreasing yield by 12.45% to 23.84%. Exogenous ABA application, especially the T3 treatment, significantly increased the grain-filling rate, extended the grain-filling duration, and enhanced grain weight, thereby mitigating drought-induced yield loss. Transcriptome sequencing generated 290.19 Gb of clean data and 23,247 differentially expressed genes (DEGs) with the highest number of DEGs in the T3 versus T1 comparison at 30 days after flowering. Subsequently, WGCNA identified ten co-expression modules, with the blue and magenta modules showing the strongest positive correlation with grain weight. Moreover, eight transcription factors (TFs) genes were identified as potential regulators of grain development under drought stress and ABA treatments.
Conclusion: Exogenous ABA application at 20 mg L-1 effectively increased the grain-filling rate and grain weight and stabilized yield under drought stress. This improvement was associated with the activation of starch and sucrose metabolism and hormone signaling pathways, potentially coordinated by key TFs including the B3, NAC, and WOX families. © 2026 Society of Chemical Industry.
(© 2026 Society of Chemical Industry.)
References: Wang HL, Zhang H, Liu JJ, Ma Q, Wu EG, Gao JF et al., Transcriptome analysis reveals the mechanism of nitrogen fertilizers in starch synthesis and quality in waxy and non‐waxy proso millet. Carbohydr Polym 323:121372 (2024).
Peng RH and Zhang BH, Foxtail millet: a new model for C4 plants. Trends Plant Sci 26:199–201 (2021).
Cao XN, Hu YL, Song J, Feng H, Wang JJ, Chen L et al., Transcriptome sequencing and metabolome analysis reveals the molecular mechanism of drought stress in millet. Int J Mol Sci 23:10792 (2022).
Liang G, Zhang WH, Du X, Guo J, Song JH, Wang XM et al., Effects of plastic‐film mulching and supplementary irrigation on yield formation and water use efficiency of hybrid millet. Acta Agronomica Sin 43:122–132 (2017).
Wang J, Sun ZX, Wang XN, Tang Y, Li XY, Ren CY et al., Transcriptome‐based analysis of key pathways relating to yield formation stage of foxtail millet under different drought stress conditions. Front Plant Sci 13:1110910 (2023).
Zhang SX, Yu Y, Song TQ, Zhang MF, Li N, Yu M et al., Genome‐wide identification of foxtail millet's TRX family and a functional analysis of SiNRX1 in response to drought and salt stresses in transgenic Arabidopsis. Front Plant Sci 13:946037 (2022).
Yang PG, Drohan PJ, Zhang XJ, Long HY, Soulis KX and Shi XR, Impacts of deficit irrigation on strawberry physiology, water productivity, quality, and yield. Sustainability 17:675 (2025).
Wang YL, Wang Y, Du JZ and Guan YA, Effects of drought stress at different periods on agronomic traits of millet. Acta Agric Boreali‐Sin 27:125–129 (2012).
Ren CY, Jiang ZQ, Su XY, Mi QC, Wang J, Li Y et al., Effects of water stress/rewatering on leaf photosynthetic characteristics and grain yield of foxtail millet. J Appl Meteorol Sci 32:456–467 (2021).
Shao J, Effects of Drought Stress at Seedling Stage and Heading Stage Onphysiological and Biochemical Characteristics and Yield of Foxtailmillet. Xinjiang Agricultural University, Urumqi, Xinjiang, China (2021).
Zhao FY, Cao HX, Ma LN, Dang XW, Wan Y, Peng ML et al., Effects of supplementary irrigation on growth, yield, and water use efficiency of millet in the loess plateau. Agric Res Arid Areas 42:44–53 (2024).
Yang BH, Yao H, Zhang JX, Li YQ, Ju YL, Zhao XF et al., Effect of regulated deficit irrigation on the content of soluble sugars, organic acids and endogenous hormones in cabernet sauvignon in the Ningxia region of China. Food Chem 312:126020 (2020).
He J, Jin Y, Palta AJ, Liu HY, Chen Z and Li FM, Exogenous ABA induces osmotic adjustment, improves leaf water relations and water use efficiency, but not yield in soybean under water stress. Agronomy 9:395 (2019).
Lu XY, Peng SF, Yin YA, Sun MH, Wu ZL, Wu XY et al., Mechanisms of exogenous ABA‐induced drought tolerance in Idesia polycarpa seedlings revealed by integrated physiological, transcriptomic, and metabolomic analyses. Ind Crop Prod 236:122030 (2025).
Deng S, Xing PP, Peng LG, Lu J, Wu YZ, Zhang YY et al., Exogenous application of abscisic acid enhanced 2‐acetyl‐1‐pyrroline biosynthesis, accumulation, and antioxidant activities in aromatic rice. BMC Plant Biol 25:302 (2025).
Luo YL, Li WQ, Huang C, Yang JH, Jin M, Chen J et al., Exogenous abscisic acid coordinating leaf senescence and transport of assimilates into wheat grains under drought stress by regulating hormones homeostasis. Crop J 9:901–914 (2021).
Cai T, Xu HC, Peng DL, Yin YP, Yang WB, Ni YL et al., Exogenous hormonal application improves grain yield of wheat by optimizing tiller productivity. Field Crop Res 155:172–183 (2014).
Travaglia C, Reinoso H, Cohen A, Luna C, Tommasino E, Castillo C et al., Exogenous ABA increases yield in field‐grown wheat with moderate water restriction. J Plant Growth Regul 29:366–374 (2010).
Chen TT, Li GY, Islam MR, Fu WM, Feng BH, Tao LX et al., Abscisic acid synergizes with sucrose to enhance grain yield and quality of rice by improving the source‐sink relationship. BMC Plant Biol 19:1–17 (2019).
Hu W, Zhang JP, Wu ZQ, Loka DA, Zhao WQ, Chen BL et al., Effects of single and combined exogenous application of abscisic acid and melatonin on cotton carbohydrate metabolism and yield under drought stress. Ind Crop Prod 176:114302 (2022).
Tang T, Xie H, Wang YX, Lv B and Liang JS, The effect of sucrose and abscisic acid interaction on sucrose synthase and its relationship to grain filling of rice (Oryza sativa L.). J Exp Bot 60:2641–2652 (2009).
Kamran M, Liu J, Lin F, Ullah A, Aqeel T, Shahzad M et al., Integrated transcriptome and metabolome association analysis reveals the complex genetic architecture of tobacco bacterial wilt resistance. Ind Crop Prod 233:121488 (2025).
Qian JJ, Zhang XP, Yan Y, Wang N, Ge WQ, Zhou Q et al., Unravelling the molecular mechanisms of abscisic acid‐mediated drought‐stress alleviation in pomegranate (Punica granatum L.). Plant Physiol Biochem 157:211–218 (2020).
Han CN, Chen GJ, Zheng DF and Feng NJ, Transcriptomic and metabolomic analyses reveal that ABA increases the salt tolerance of rice significantly correlated with jasmonic acid biosynthesis and flavonoid biosynthesis. Sci Rep 13:20365 (2023).
Urbutis M, Vaseva II, Simova‐Stoilova L, Todorova D, Pukalskas A and Samuolience G, Drought protective effects of exogenous ABA and kinetin on lettuce: sugar content, antioxidant enzyme activity, and productivity. Plants 13:1641 (2024).
Xiang L, Wang D, Gu SB, White PJ, Han K, Zhou J et al., Effect of supplemental irrigation on the relationships between leaf ABA concentrations, tiller development and photosynthate accumulation and remobilization in winter wheat. Plant Growth Regul 79:331–343 (2016).
Wang JW, Li Y and Niu WQ, Deficit alternate drip irrigation increased root‐soil‐plant interaction, tomato yield, and quality. Int J Environ Res Public Health 17:781 (2020).
Jiang Z, Piao L, Guo D, Zhu H, Wang S, Zhu H et al., Regulation of maize kernel carbohydrate metabolism by abscisic acid applied at the grain‐filling stage at low soil water potential. Sustainability 13:3125 (2020).
Hong ZQ, Zhang ZZ, Wang L, Su M, Zhou T, Li FG et al., Controlled‐release and conventional urea combinations enhance post‐flowering photosynthesis and grain filling of spring maize in the Ningnan mountainous area. J Plant Nutr 31:1983–1997 (2025).
Zhang YX, Bao PX, Li WJ, Zhang X, Zang ZY, Zhang WL et al., Effects of grain‐filling characteristics of different varieties of setaria italica var. germanica on yield formation in jilin province. Seed 44:167–173 (2025).
Dong Q, Zhang Y, Zhu DY, Peng YL, Liu RT, Ya MF et al., Effect of different nitrogen fertilizer treatments on grain filling characteristics of black highland barley. Journal of Triticeae Crops 43:940–946 (2023).
Li ML, Liang ZY, Zhang HY, Yang Y, He DH, Luo T et al., Effects of exogenous α‐ketoglutaric acid spraying after anthesis on photosynthetic characteristics, grain filling and yield of wheat. J Triticeae Crops 45:1394–1402 (2025).
Livak KJ and Schmittgen TD, Analysis of relative gene expression data using real‐time quantitative PCR and the 2(‐Delta Delta C(T)). Method Methods (San Diego, Calif) 25:402–408 (2001).
Shi C, Qiu T, Zhang YY, Ma Y, Li XR, Dong SQ et al., Effects of different preceding crops on soil nutrients and foxtail millet productivity and quality. Front Plant Sci 15:1477756 (2024).
Gao XL, Ma JJ, Jia YR, Liu EK and Song LL, The effects of deficit irrigation scheduling on water consumption and water use efficiency of millet in the northern shanxi province. J Irrig Drain 40:40–47 (2021).
Liu J, Determination and Genome‐Wide Association Study of Wheat Grain Filling Rate. Northwest A&F University, Yangling, Shaanxi, China (2024).
Zhu GH, Ye NH, Yang JC, Peng XX and Zhang JH, 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 (2021).
Zhuang TX, Ata‐U‐Karim ST, Zhao B, Liu XJ, Tian YC, Zhu Y et al., Investigating the impacts of different degrees of deficit irrigation and nitrogen interactions on assimilate translocation, yield, and resource use efficiencies in winter wheat. Agric Water Manag 304:109089 (2024).
Yang DQ, Li YL, Ni YL, Yin YP, Yang WB, Cui ZY et al., Effects of exogenous ABA and 6‐BA on protein content and grain filling process in different types of stay‐green wheat. Acta Agron Sin 40:301–312 (2014).
Yu T, Xin YN and Wang J, Effects of exogenous abscisic acid on grain filling characteristics, starch accumulation, and endogenous hormones in maize under early post‐anthesis high temperature stress. Chin J Appl Ecol 35:2715–2724 (2024).
Yu T, Xin Y and Liu P, Exogenous abscisic acid (ABA) improves the filling process of maize grains at different ear positions by promoting starch accumulation and regulating hormone levels under high planting density. BMC Plant Biol 24:80 (2024).
Hussain S, Saleem MF, Javaid Iqbal J, Ibrahim M, Atta S, Ahmed T et al., Exogenous application of abscisic acid may improve the growth and yield of sunflower hybrids under drought. Pak J Agric Sci 51:49–58 (2014).
Dash GK, Geda CK, Pati S, Sabrinathan S, Saho SK, Purkaystha S et al., Exogenously applied ABA modulated enzymatic changes conferring drought tolerance in rice varieties. Cereal Res Commun 2024:1–13 (2024).
Gu JM, Liu PK, Nie WT, Wang ZJ, Cui XY, Fu HD et al., Abscisic acid alleviates photosynthetic damage in the tomato ABA‐deficient mutant sitiens and protects photosystem II from damage via the WRKY22–PsbA complex under low‐temperature stress. J Integr Agric 24:546–563 (2025).
Rezaul M, Feng BH, Chen TT, Fu WM, Zhang CX, Tao LX et al., Abscisic acid prevents pollen abortion under high‐temperature stress by mediating sugar metabolism in rice spikelets. Physiol Plant 165:644–663 (2019).
Dewdney J and McWha JA, Abscisic acid and the movement of photosynthetic assimilates towards developing wheat (Triticum aestivum L.) grain. Zi für Pflanzenphysiol 92:183–186 (1979).
Zhang WJ, Zhao Y, Li LY, Xu X, Yang L, Luo Z et al., The effects of short‐term exposure to low temperatures during the booting stage on starch synthesis and yields in wheat grain. Front Plant Sci 12:684784 (2021).
Sun LY, Zu X, Cui SK, Zhu MW, Huang T, Duan Y et al., Postharvest water loss‐induced transcriptomic reprogramming modulates non‐structural carbohydrate remobilization in Quercus texana kernels. Ind Crop Prod 227:120768 (2025).
Yang TY, Li HR and Li LW, Genome‐wide characterization and expression analysis of α‐amylase and β‐amylase genes underlying drought tolerance in cassava. BMC Genomics 24:190 (2023).
Xiao QL, Liu TT, Ling M, Ma QN, Cao W, Xing FY et al., Genome‐wide identification of DOF gene family and the mechanism dissection of SbDof21 regulating starch biosynthesis in sorghum. Int J Mol Sci 23:12152 (2022).
Gao Y, Jing MY, Zhang M, Zhang Z, Liu YQ, Wang ZM et al., Transcriptomic and metabolomic analysis of changes in grain weight potential induced by water stress in wheat. J Integr Agric 23:3706–3722 (2024).
Pattanagul W, Exogenous abscisic acid enhances sugar accumulation in Rice (Oryza sativa L.) under drought stress. Asian J Plant Sci 10:212–219 (2011).
Gai ZS, Wang Y, Ding YQ, Qian WJ, Qiu C, Xie H et al., Exogenous abscisic acid induces the lipid and flavonoid metabolism of tea plants under drought stress. Sci Rep 10:12275 (2020).
Gowsiga S, Vijayalakshmi D, Deepika J, Arunkumar P and Sivakumar R, Stomatal traits, hormone and metabolite profiles governing drought and high temperature stress tolerance in foxtail millet. Russian J Plant Physiol 72:42 (2025).
Jiang ZZ, Li YP, Gai PZ, Gao JZ and Xu LJ, Exogenously applied ABA alleviates dysplasia of maize (Zea mays L.) ear under drought stress by altering photosynthesis and sucrose transport. Plant Signal Behav 20:2462497 (2025).
Zhang QW, Song XM and Bartels D, Sugar metabolism in the desiccation tolerant grass Oropetium thomaeum in response to environmental stresses. Plant Sci 270:30–36 (2018).
Xiong YL, Song XY, Mehra P, Yu SH, Li QY, Tashenmaimaiti D et al., ABA‐auxin cascade regulates crop root angle in response to drought. Curr Biol 35:542–553 (2025).
Norma F, Takuya Y and Alisdair RF, Role of Raf‐like kinases in SnRK2 activation and osmotic stress response in plants. Nat Commun 11:6184 (2020).
Abhilasha A and Roy Choudhury S, Molecular and physiological perspectives of abscisic acid mediated drought adjustment strategies. Plants 10:2769 (2021).
Hu W, Yan Y and He YZ, Cloning and expression analysis of TaPP2C59 gene in wheat. J Triticeae Crops 34:1334 (2024).
Yu TF, Liu Y, Fu JD, Ma J, Fang ZW, Chen J et al., The NF‐Y‐PYR module integrates the abscisic acid signal pathway to regulate plant stress tolerance. Plant Biotechnol J 19:2589–2605 (2021).
Teng ZN, Yu HH, Wang GQ, Meng S, Liu BH, Yi YK et al., Synergistic interaction between ABA and IAA due to moderate soil drying promotes grain filling of inferior spikelets in rice. Plant J: Cell Mol Biol 109:1457–1472 (2021).
Xing LP, Di ZC, Yang WW, Liu JQ, Li MN, Wang XJ et al., Overexpression of ERF1‐V from Haynaldia villosa can enhance the resistance of wheat to powdery mildew and increase the tolerance to salt and drought stresses. Front Plant Sci 8:1948 (2017).
Yan D, Li CN, Mao XG, Wang JY, Long L, Yang JW et al., TaERF73 is associated with root depth, thousand‐grain weight and plant height in wheat over a range of environmental conditions. Food Energy Secur 11:325 (2021).
Bu YN, Song TQ, Zhou JF, Lei XP, Chang XL, Li SY et al., The wheat HSF transcription factor TaHSFC3B confers drought tolerance through ROS scavenging and ABA pathway in transgenic Arabidopsis and wheat (Triticum aestivum L.). Planta 262:1–24 (2025).
Liu W, Zhang YW, Wang YB, Li W, Xu R, Wang CJ et al., Screening of soybean drought responsive gras genes and bioinformatics and adversity stress expression analysis on gmgras27. Soybean Sci 41:36–42 (2022).
Cui ZB, Wang XW, Dai YD, Li YY, Ban YJ, Tian WJ et al., Transcription factor OsNF‐YC1 regulates grain size by coordinating the transcriptional activation of OsMADS1 in Oryza sativa L. Plant J: Cell Mol Biol 119:1465–1480 (2024).
Wang YW, Liao YR, Quan CQ, Li YQ, Yang SJ, Ma C et al., C2H2‐type zinc finger OsZFP15 accelerates seed germination and confers salinity and drought tolerance of rice seedling through ABA catabolism. Environ Exp Bot 199:104873 (2022).
Huang XX, Huang LP, Zhao XX, Jia J, Zhang G, Zhang MY et al., A J‐protein OsDjC46 interacts with ZFP36 to participate in ABA‐mediated antioxidant defense in Rice. Antioxidants (Basel) 11:207 (2022).
Tong ZH, Zhang YL, Wang LL, Li AR, Wang WQ, Wang PP et al., Genome wide identification and expression pattern analysis of B3 gene family in maize. Pratacultural Sci 40:2556–2570 (2023).
Yang T, Guo LX, Ji C, Wang HH, Wang JC, Zheng XX et al., The B3 domain‐containing transcription factor ZmABI19 coordinates expression of key factors required for maize seed development and grain filling. Plant Cell 33:104–128 (2021).
Pan LY, Wang YJ, Li HJ, Hou F, Li J, Li LL et al., Screening regulatory genes related to wheat grain protein accumulation based on transcriptome and WGCNA analysis. Sci Agric Sin 58:1065–1082 (2025).
Li JH, Xie LN, Tian XL, Liu SY, Xu DG, Jin H et al., TaNAC100 acts as an integrator of seed protein and starch synthesis conferring pleiotropic effects on agronomic traits in wheat. Plant J: Cell Mol Biol 108:829–840 (2021).
Liu HX, Li T, Hou J, Yin XT, Wang YQ, Si XM et al., TaWUS‐like‐5D affects grain weight and filling by inhibiting the expression of sucrose and trehalose metabolism‐related genes in wheat grain endosperm. Plant Biotechnol J 23:2018–2033 (2025).
Grant Information: the National Key Research and Development Plan of China Programs (2021YFD1900705) and the National Key Research and Development Program of China (2022YFD1901602). National Natural Science Foundation of China (32301315), the Innovation Capability Support Program of Shaanxi Province (2023KJXX-072 and 2024CX-GXPT-03)
Contributed Indexing: Keywords: WGCNA; drought stress; exogenous ABA application; foxtail millet; transcriptomic analysis
Substance Nomenclature: 72S9A8J5GW (Abscisic Acid)
0 (Plant Proteins)
0 (Plant Growth Regulators)
Entry Date(s): Date Created: 20260508 Date Completed: 20260708 Latest Revision: 20260708
Update Code: 20260709
DOI: 10.1002/jsfa.70704
PMID: 42101208
Βάση Δεδομένων: MEDLINE
Περιγραφή
ISSN:1097-0010
DOI:10.1002/jsfa.70704