Journal of Plant Growth Regulation

, Volume 38, Issue 4, pp 1467–1480 | Cite as

Effects of Plant Growth Regulators on Seed Filling, Endogenous Hormone Contents and Maize Production in Semiarid Regions

  • Irshad AhmadEmail author
  • Muhammad Kamran
  • Xiangping Meng
  • Shahzad Ali
  • Bayasgalan Bilegjargal
  • Tie Cai
  • Tiening Liu
  • Qingfang HanEmail author


In this study, we determined whether the application of uniconazole alone or combined with ethephon could enhance the seed-filling rates in maize by regulating the endogenous hormone contents. Uniconazole was applied to the foliage at concentrations of 0 (CK), 25 (U25), 50 (U50) and 75 (U75) mg L−1 at the 12-leaf stage. In addition, uniconazole was applied to the foliage at the 12-leaf stage and ethephon at 10 days after silking stage at concentrations of 0 (CK), 25 mg L−1 uniconazole + 100 mg L−1 ethephon (U25 + E100), 50 mg L−1 uniconazole + 200 mg L−1 ethephon (U50 + E200) and 75 mg L−1 uniconazole + 300 mg L−1 ethephon (U75 + E300). Uniconazole applied alone or in combination with ethephon significantly improved ear characters and grain yield. Uniconazole applied alone or combination with ethephon significantly improved the dry matter accumulation in seeds and seed-filling rates. Uniconazole significantly increased the abscisic acid (ABA) and zeatin (Z) + zeatin riboside (ZR) contents of seeds, but reduced the gibberellic acid (GA) contents. The application of uniconazole combined with ethephon decreased the ABA, Z + ZR and GA contents in seeds. The ABA and Z + ZR contents were significantly positively correlated, whereas the GA content was negatively correlated with the maximum seed weight, maximum seed-filling rate and mean seed-filling rate. The application of uniconazole alone significantly improved the seed-filling rates in maize by regulating the endogenous hormone contents. Thus, we conclude that uniconazole application at 50 mg L−1 in the 12-leaf stage can enhance the maize production.


Ethephon Hormones content Maize Semiarid regions Seed-filling Uniconazole 



The authors extend their sincere thanks to the editors of this journal and the anonymous reviewers for their valuable comments and suggestions that have significantly improved the manuscript. We are also grateful to Ding Ruixia, Nie Junfeng and Yang Baoping for help during experimental period.

Author Contributions

QH, LT and TC conceived and designed the research. IA performed research. MK, XM and SA contributed in the field experiments. IA wrote the manuscript. BB helped in English revision.


This study was supported by funding from High Technology Research and Development Program of China (863 Program, No. 2013AA102902), the National Natural Science Foundation of China (No. 31601256), the special fund for Agro-scientific Research in the Public Interest (201303104), the 111 Project of Chinese Education Ministry (B12007).

Compliance with Ethical Standards

Conflict of interest

The authors have no conflict of interest.


  1. Ahmad I, Kamran M, Ali S, Bilegjargal B, Cai T, Ahmad S, Meng XP, Su W, Liu T, Han QF (2018a) Uniconazole application strategies to improve lignin biosynthesis, lodging resistance and production of maize in semiarid regions. Field Crops Res 222:66–77Google Scholar
  2. Ahmad I, Kamran M, Su W, Haiqi W, Ali S, Bilegjargal B, Ahmad S, Liu T, Cai T, Han QF (2018b) Application of uniconazole improves photosynthetic efficiency of maize by enhancing the antioxidant defense mechanism and delaying leaf senescence in semiarid regions. J Plant Growth Regul. CrossRefGoogle Scholar
  3. Alferez F, Pozo L, Burns JK (2006) Physiological changes associated with senescence and abscission in mature citrus fruit induced by 5-chloro-3-methyl-4-nitro-1H-pyrazole and ethephon application. Physiol Plantarum 127:66–73Google Scholar
  4. Ali S, Jan A, Zhang P, Khan MN, Cai T, Wei T, Ren X, Jia Q, Han Q, Jia Z (2016) Effects of ridge-covering mulches on soil water storage and maize production under simulated rainfall in semiarid regions of China. Agric Water Manag 178:1–11Google Scholar
  5. Ali S, Xu YY, Ahmad I, Jia QM, Fangyuan H, Daur I, Wei T, Cai T, Ren XL, Zhang P, Jia ZK (2018) The ridge furrow cropping technique indirectly improves seed filling endogenous hormonal changes and winter wheat production under simulated rainfall conditions. Agric Water Manag 204:138–148Google Scholar
  6. Alves AAC, Setter TL (2004) Abscisic acid accumulation and osmotic adjustment in cassava under water deficit. Environ Exp Bot 51:259–271Google Scholar
  7. Bakheta MA, Hussein MM (2014) Uniconazole effect on endogenous hormones, proteins and proline contents of barley plants (Hordium vulgare) under salinity stress (NaCl). Nusantara Biosci 6:2087–3948Google Scholar
  8. Bidadi H, Yamaguchi S, Asahina M, Satoh S (2010) Effects of shoot-applied gibberellin/gibberellin-biosynthesis inhibitors on root growth and expression of gibberellin biosynthesis genes in Arabidopsis thaliana. Plant Root 4:4–11Google Scholar
  9. Burden RS, Carter GA, Clark T, Cooke DT, Croker SJ, Deas AHB, Hedden P, James CS, Lenton JR (1987) Comparative activity of the enantiomers of triadimenol and paclobutrazol as inhibitors of fungal growth and plant sterol and gibberellin biosynthesis. Pestic Sci 21:253–267Google Scholar
  10. Chen HJ, Tsai YJ, Chen WS, Huang GJ, Huang SS, Lin YH (2010) Ethephon-mediated effects on leaf senescence are affected by reduced glutathione and EGTA in sweet potato detached leaves. Bot Stud 51:171–181Google Scholar
  11. Chen K, Ali S, Chen Y, Sohail A, Jan A, Fahad S (2018) Effect of ridge-covering mulching materials on hormonal changes, antioxidative enzyme activities and production of maize in semi-arid regions of China. Agric Water Manag 204:281–291Google Scholar
  12. Costa ML, Civello PM, Chaves AR, Martinez GA (2005) Effect of ethephon and 6-benzylaminopurine on chlorophyll degrading enzymes and a peroxidase-linked chlorophyll bleaching during post-harvest senescence of broccoli (Brassica oleracea L.) at 20 degrees C. Postharvest Biol Tech 35:191–199Google Scholar
  13. Duan L, Guan C, Li J, Eneji AE, Li Z, Zhai Z (2008) Compensative effects of chemical regulation with uniconazole on physiological damages caused by water deficiency during the grain filling stage of wheat. J Agron Crop Sci 194:9–14Google Scholar
  14. Fang X, Liu X, Zhang Y, Huang K, Zhang Y, Li Y, Nie J, She H, Ruan R, Yuan X, Yi Z (2018) Effects of uniconazole or gibberellic acid application on the lignin metabolism in relation to lodging resistance of culm in common buckwheat (Fagopyrum esculentum M.). J Agron Crop Sci 204:414–423Google Scholar
  15. Fletcher RA, Kallidumbil V, Steele P (1982) An improved bioassay for cytokinins using cucumber cotyledons. Plant Physiol 69:675–677PubMedPubMedCentralGoogle Scholar
  16. Grassini P, Yang H, Irmak S, Thorburn J, Burr C, Cassman KG (2011) High-yield irrigated maize in the Western U.S. Corn Belt: II. Irrigation management and crop water productivity. Field Crops Res 120:133–141Google Scholar
  17. Han H, Yang WY (2009) Influence of uniconazole and plant density on nitrogen content and grain quality in winter wheat in South China. Plant Soil Environ 55:159–166Google Scholar
  18. Han Y, Gao Y, Shi Y, Du J, Zheng D, Liu G (2017) Genome-wide transcriptome profiling reveals the mechanism of the effects of uniconazole on root development in Glycine Max. J Plant Biol 60:387–403Google Scholar
  19. Huang MJ, Fang Y, Liu Y, Jin Y, Sun JL, Tao X, Ma XR, He KZ, Zhao H (2015) Using proteomic analysis to investigate uniconazole-induced phytohormone variation and starch accumulation in duckweed (Landoltia punctata). BMC Biotechnol 15:81PubMedPubMedCentralGoogle Scholar
  20. Hussein MM, Bakheta MA, Zaki SS (2014) Influence of uniconazole on growth characters, photosynthetic pigments, total carbohydrates and total soluble sugars of Hordium vulgare L. plants grown under salinity stress. Int J Sci Res 3:2208–2213Google Scholar
  21. Ijaz M, Honermeier B (2012) Effect of triazole and strobilurin fungicides on seed yield formation and grain quality of winter rapeseed (Brassica napus L.). Field Crops Res 130:80–86Google Scholar
  22. Izumi K, Kamiya Y, Sakurai A, Oshio H, Takahashi N (1985) Studies of sites of action of a new plant growth retardant (E)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol (S-3307) and comparative effects of its stereoisomers in a cell-free system from Cucurbita maxima. Plant Cell Physiol 26:821–827Google Scholar
  23. Izumi K, Nakagawa S, Kobayashi M, Oshio H, Sakurai A, Takahashi N (1988) Levels of IAA, cytokinins, ABA and ethylene in rice plants as affected by a gibberellins biosynthesis inhibitor uniconazole P. Plant Cell Physiol 29:97–104Google Scholar
  24. Kamran M, Cui W, Ahmad I, Meng XP, Zhang X, Su W, Chen J, Ahmad S, Fahad S, Han QF, Tiening L (2018a) Effect of paclobutrazol, a potential growth regulator on stalk mechanical strength, lignin accumulation and its relation with lodging resistance of maize. Plant Growth Regul 84:317–332Google Scholar
  25. Kamran M, Ahmad I, Wang HQ, Wu XR, Xu J, Tiening L, Ding RX, Han QF (2018b) Mepiquat chloride application increases lodging resistance of maize by enhancing stem physical strength and lignin biosynthesis. Field Crops Res 224:148–159Google Scholar
  26. Kamran M, Su W, Ahmad I, Xiangping M, Wenwen C, Xudong Z, Siwei M, Khan A, Qingfang H, Tiening L (2018c) Application of paclobutrazol affect maize grain yield by regulating root morphological and physiological characteristics under a semi-arid region. Sci Rep 8:4818PubMedPubMedCentralGoogle Scholar
  27. Lee KH, Piao HL, Kim HY, Choi SM, Jiang F, Hartung W, Hwang I, Kwak JM, Lee IJ, Hwang I (2006) Activation of glucosidase via stress-induced polymerization rapidly increases active pools of abscisic acid. Cell 26:1109–1120Google Scholar
  28. Lefebvre V, North H, Frey A, Sotta B, Seo M, Okamoto M, Nambara E, Marion-Poll A (2006) Functional analysis of Arabidopsis NCED6 and NCED9 genes indicates that ABA synthesized in the endosperm is involved in the induction of seed dormancy. Plant J 45:309–319PubMedGoogle Scholar
  29. Liu Y, Sui YW, Gu DD, Wen XX, Chen Y, Li CJ, Liao YC (2013a) Effects of conservation tillage on grain filling and hormonal changes in wheat under simulated rainfall conditions. Field Crops Res 144:43–51Google Scholar
  30. Liu Y, Han J, Wen XX, Wu W, Guo Q, Zeng AL, Liao YC (2013b) Effect of plasticcovered ridge and furrow planting on grain filling of winter wheat is related to changes in endogenous hormones. J Integr Agric 12:1771–1782Google Scholar
  31. Liu Y, Han J, Liu DD, Gu DD, Wang YP, Liao YC, Wen XX (2015a) Effect of plastic film mulching on the grain filling and hormonal changes of maize under different irrigation conditions. PLoS ONE 10:e0122791PubMedPubMedCentralGoogle Scholar
  32. Liu Y, Fang Y, Huang M, Jin Y, Sun J, Tao X, Zhang G, He K, Zhao Y, Zhao H (2015b) Uniconazole-induced starch accumulation in the bioenergy crop duckweed (Landoltiapunctata) II: transcriptome alterations of pathways involved in carbohydrate metabolism and endogenous hormone crosstalk. Biotechnol Biofuels 8:1–12Google Scholar
  33. Liu Y, Fang Y, Huang M, Jin Y, Sun J, Tao X, Zhang G, He K, Zhao Y, Zhao H (2015c) Uniconazole-induced starch accumulation in the bioenergy crop duckweed (Landoltia punctata) I: transcriptome analysis of the effects of uniconazole on chlorophyll and endogenous hormone biosynthesis. Biotech Biofuels 8:57Google Scholar
  34. Liu Y, Liang HY, Lv XK, Liu DD, Wen XX, Liao YC (2016) Effect of polyamines on the grain filling of wheat under drought stress. Plant Physiol Biochem 100:113–129PubMedGoogle Scholar
  35. Liu Y, Chen XY, Wang XH, Fang Y, Huang MJ, Guo L, Zhang Y, Zhao H (2018) Improving biomass and starch accumulation of bioenergy crop duckweed (Landoltia punctata) by abscisic acid application. Sci Rep 8:9544PubMedPubMedCentralGoogle Scholar
  36. Lv XK, Li T, Wen XX, Liao YC, Liu Y (2017a) Effect of potassium foliage application post-anthesis on grain filling of wheat under drought stress. Field Crops Res 206:95–105Google Scholar
  37. Lv XK, Han J, Liao YC, Liu Y (2017b) Effect of phosphorus and potassium foliage application post-anthesis on grain filling and hormonal changes of wheat. Field Crops Res 214:83–93Google Scholar
  38. Mares DJ, Marschner H, Krauss A (1981) Effect of gibberellic-acid on growth and carbohydrate-metabolism of developing tubers of potato (solanum-tuberosum). Plant Physiol 52:267–274Google Scholar
  39. Nambara E, Marion-Poll A (2005) Abscisic acid biosynthesis and catabolism. Ann Rev Plant Biol 56:165–185Google Scholar
  40. National Bureau of Statistics of China (NBSC) (2012) China statistics yearbook. China Statistics Press, Beijing.
  41. Parry M, Lowe J, Hanson C (2009) Overshoot, adapt and recover. Nature 458:1102–1103PubMedGoogle Scholar
  42. Piotrowska A, Bajguz A (2011) Conjugates of abscisic acid, brassinosteroids, ethylene, gibberellins, and jasmonates. Phytochem 72:2097–2112Google Scholar
  43. Qin SJ, Zhang ZZ, Ning TY, Ren SZ, Su LC, Li ZJ (2013) Abscisic acid and aldehyde oxidase activity in maize ear leaf and grain relative topost-flowering photosynthetic capacity and grain-filling rate under different water/nitrogen treatments. Plant Physiol Biochem 70:69–80PubMedGoogle Scholar
  44. Rahim AAOS, Elamin OM, Bangerth FK (2011) Effects of paclobutrazol (PBZ) on floral induction and associated hormonal and metabolic changes of biennially bearing mango (Mangifera indica L.) cultivars during off year. J Agric Biol Sci 6:55–67Google Scholar
  45. Ren X, Chen X, Jia Z (2010) Effect of rainfall collecting with ridge and furrow on soil moisture and root growth of corn in semiarid Northwest China. J Agron Crop Sci 196:109–122Google Scholar
  46. Rentzsch S, Podzimska D, Voegele A, Imbeck M, Mueller K, Linkies A, Leubner-Metzger G (2012) Dose- and tissue-specific interaction of monoterpenes with the gibberellin-mediated release of potato tuber bud dormancy, sprout growth and induction of alpha-amylases and beta-amylases. Planta 235:137–151PubMedGoogle Scholar
  47. Richards FJ (1959) A flexible growth function for empirical use. J Exp Bot 10:290–301Google Scholar
  48. Saito S, Okamoto M, Shinoda S, Kushiro T, Koshiba T, Kamiya Y, Hirai N, Todoroki Y, Sakata K, Nambara E, Mizutani M (2006) A plant growth retardant, uniconazole, is a potent inhibitor of ABA catabolism in arabidopsis. Biosci Biotechnol Biochem 70:1731–1739PubMedGoogle Scholar
  49. Soumya PR, Kumar P, Pal M (2017) Paclobutrazol: a novel plant growth regulator and multi-stress ameliorant. Ind J Plant Physiol 22:267–278Google Scholar
  50. Tripathi SC, Sayre KD, Kaul JN, Narang RS (2003) Growth and morphology of spring wheat (Triticum aestivum L.) culms and their association with lodging: effects of genotypes, N levels and ethephon. Field Crops Res 84:271–290Google Scholar
  51. Verslues PE, Zhu J-K (2005) Before and beyond ABA: upstream sensing and internal signals that determine ABA accumulation and response under abiotic stress. Biochem Soc Trans 33:375–379PubMedGoogle Scholar
  52. Wang XC, Yang WY, Chen G, Qian-Liang LI, Wang XB (2009) Effects of uniconazole on leaf senescence and yield of maize sprayed at late growth stage. J Maize Sci 17:86–88Google Scholar
  53. Wang HY, Zhang MC, Chen XW, Niu XX, Su H, Dong XH (2013) Effects of ethephon on dry matter accumulation in summer corn seeds and related physiological mechanism inquiry. J Maize Sci 21:57–61Google Scholar
  54. Wang C, Hu D, Liu X, She H, Ruan R, Yang H, Yi Z, Wu D (2015) Effects of uniconazole on the lignin metabolism and lodging resistance of culm in common buckwheat (Fagopyrum esculentum M.). Field Crops Res 180:46–53Google Scholar
  55. Xu YJ, Gu DJ, Zhang BB, Zhang H, Wang ZQ, Yang JC (2013) Hormone contents in kernels at different positions on an ear and their relationship with endosperm development and kernel filling in maize. Acta Agron Sin 39:1452–1461Google Scholar
  56. Yang J, Zhang J (2006) Grain filling of cereals under soil drying. New Phytol 169:223–236PubMedGoogle Scholar
  57. Yang J, Peng S, Visperas RM, Sanico AL, Zhu Q, Gu S (2000) Grain filling pattern and cytokinin content in the grains and roots of rice plants. Plant Growth Regul 30:261–270Google Scholar
  58. Yang J, Zhang J, Wang Z, Zhu Q, Wang W (2001) Hormonal changes in the grains of rice subjected to water stress during grain filling. Plant Physiol 127:315–323PubMedPubMedCentralGoogle Scholar
  59. Yang WY, Xiang ZF, Ren WJ, Wang XC (2005) Effect of S-3307 on nitrogen metabolism and grain protein content in rice. Chin J Rice Sci 19:63–67Google Scholar
  60. Yang J, Zhang J, Wang Z, Liu K, Wang P (2006) Post-anthesis development of inferior and superior spikelets in rice in relation to abscisic acid and ethylene. J Exp Bot 57:149–160PubMedGoogle Scholar
  61. Ye DL, Zhang YS, Alkaisi MM, Duan LS, Zhang MC, LI ZH (2016) Ethephon improved stalk strength associated with summer maize adaptations to environments differing in nitrogen availability in the North China Plain. J Agric Sci 154:960–977Google Scholar
  62. Zhang MC, Duan LS, Tian XL, He ZP, Li JM, Wang BM, Li ZH (2007) Uniconazole-induced tolerance of soybean to water deficit stress in relation to changes in photosynthesis, hormones and antioxidant system. J Plant Physiol 164:709–717Google Scholar
  63. Zhang H, Tan G, Yang L, Yang J, Zhang J, Zhao B (2009) Hormones in the grains and roots in relation to post-anthesis development of inferior and superior spikelets in japonica/indica hybrid rice. Plant Physiol Biochem 47:195–204PubMedGoogle Scholar
  64. Zhang H, Chen TT, Wang ZQ, Yang JC, Zhang JH (2010) Involvement of cytokinins in the grain filling of rice under alternate wetting and drying irrigation. J Exp Bot 61:3719–3733PubMedGoogle Scholar
  65. Zhang Z, Chen J, Lin S, Li Z, Cheng R, Fang C, Chen H, Lin W (2012) Proteomic and phosphorproteomic determination of ABA’s effects on grain-filling of Oryza sativa L. inferior spikelets. Plant Sci 185:259–273PubMedGoogle Scholar
  66. Zhou W, Leul M (1998) Uniconazole-induced alleviation of freezing injury in relation to changes in hormonal balance, enzyme activities and lipid peroxidation in winter rape. Plant Growth Regul 26:41–47Google Scholar
  67. Zhou W, Leul M (1999) Uniconazole-induced tolerance of rape plants to heat stress in relation to changes in hormonal levels, enzyme activities and lipid peroxidation. Plant Growth Regul 27:99–104Google Scholar
  68. Zhu QS, Cao X, Luo Y (1988) Growth analysis in the process of grain filling in rice. Acta Agron Sin 14:182–192Google Scholar

Copyright information

© Springer Science+Business Media, LLC, part of Springer Nature 2019

Authors and Affiliations

  • Irshad Ahmad
    • 1
    • 2
    Email author
  • Muhammad Kamran
    • 1
    • 2
  • Xiangping Meng
    • 1
    • 2
  • Shahzad Ali
    • 1
    • 2
  • Bayasgalan Bilegjargal
    • 3
  • Tie Cai
    • 1
    • 2
  • Tiening Liu
    • 1
    • 2
  • Qingfang Han
    • 1
    • 2
    Email author
  1. 1.College of Agronomy, Key Laboratory of Crop Physio-ecology and Tillage Science in North-western Loess Plateau, Ministry of AgricultureNorthwest A&F UniversityYanglingChina
  2. 2.Key Laboratory of Agricultural Soil and Water Engineering in Arid and Semiarid Areas, Ministry of Education, Institute of Water Saving Agriculture in Arid Areas of ChinaNorthwest A&F UniversityYanglingChina
  3. 3.College of Economics and ManagementNorthwest A&F UniversityYanglingChina

Personalised recommendations