Bian D H, Jia G P, Cai L J, Ma Z Y, Eneji A E, Cui Y H. 2016. Effects of tillage practices on root characteristics and root lodging resistance of maize. Field Crops Research, 185, 89–96.

Brune P F, Baumgarten A, McKay S J, Technow F, Podhiny J J. 2018. A biomechanical model for maize root lodging. Plant and Soil, 422, 397–408.

Buer C S, Sukumar P, Muday G K. 2006. Ethylene modulates flavonoid accumulation and gravitropic responses in roots of Arabidopsis. Plant Physiology, 140, 1384–1396.

Chen P P, Gu X B, Li Y N, Qiao L R, Li Y P, Fang H, Yin M H, Zhou C M. 2022. Effects of residual film on maize root distribution, yield and water use efficiency in Northwest China. Agricultural Water Management, 260, 107289.

Ciampitti I A, Vyn T J. 2012. Physiological perspectives of changes over time in maize yield dependency on nitrogen uptake and associated nitrogen efficiencies: A review. Field Crops Research, 133, 48–67.

Clark D G, Gubrium E K, Barrett J E, Nell T A, Klee H J. 1999. Root formation in ethylene-insensitive plants. Plant Physiology, 121, 53–60.

Dathe A, Postma J A, Postma-Blaauw M B, Lynch J P. 2016. Impact of axial root growth angles on nitrogen acquisition in maize depends on environmental conditions. Annals of Botany, 118, 401–414.

FAO (Food and Agriculture Organization of the United Nations). 2021. Food and agriculture organization of the united nations agriculture databases (FAO). [2024-12-16]. https://faostat. fao. org/

Flint-Garcia S A, Jampatong C, Darrah L L, McMullen M D. 2003. Quantitative trait locus analysis of stalk strength in four maize populations. Crop Science, 43, 13–22.

Fouéré A, Pellerin S, Duparque A. 1995. A portable electronic device for evaluating root lodging resistance in maize. Agronomy Journal, 87, 1020–1024.

Gao J, Liu Z, Wang P, Huang S B. 2024. Drip irrigation coupled with appropriate N input increased maize (Zea mays L.) yield and lodging resistance via optimizing root and stem trait. European Journal of Agronomy, 160, 127298.

Gong F P, Wu X L, Zhang H Y, Chen Y H, Wang W. 2015. Making better maize plants for sustainable grain production in a changing climate. Frontiers in Plant Science, 6, 805.

Grassini P, Thorburn J, Burr C, Cassman K G. 2011. High-yield irrigated maize in the Western U.S. Corn Belt: I. On-farm yield, yield potential, and impact of agronomic practices. Field Crops Research, 120, 142–150.

Hay R K M. 1995. Harvest index: A review of its use in plant breeding and crop physiology. Annals of Applied Biology, 126, 197–216.

Hébert Y, Guingo E, Loudet O. 2001. The response of root/shoot partitioning and root morphology to light reduction in maize genotypes. Crop Science, 41, 363–371.

Hecht V L, Temperton V M, Nagel K A, Rascher U, Postma J A. 2016. Sowing density: A neglected factor fundamentally affecting root distribution and biomass allocation of field grown spring barley (Hordeum vulgare L.). Frontiers in Plant Science, 7, 944.

Den Herder G, Van Isterdael G, Beeckman T, De Smet I. 2010. The roots of a new green revolution. Trends in Plant Science, 15, 600–607.

Hetz W, Hochholdinger F, Schwall M, Feix G. 1996. Isolation and characterization of rtcs, a maize mutant deficient in the formation of nodal roots. The Plant Journal, 10, 845–857.

Hirel B, Le Gouis J, Ney B, Gallais A. 2007. The challenge of improving nitrogen use efficiency in crop plants: Towards a more central role for genetic variability and quantitative genetics within integrated approaches. Journal of Experimental Botany, 58, 2369–2387.

Hochholdinger F. 2016. Untapping root system architecture for crop improvement. Journal of Experimental Botany, 67, 4431–4433. 

Hochholdinger F, Yu P, Marcon C. 2018. Genetic control of root system development in maize. Trends in Plant Science, 23, 79–88.

Hostetler A N, Erndwein L, Ganji E, Reneau J W, Killian M L, Sparks E E. 2022a. Maize brace root mechanics vary by whorl, genotype and reproductive stage. Annals of Botany, 129, 657–668.

Hostetler A N, Erndwein L, Reneau J W, Stager A, Tanner H G, Cook D, Sparks E E. 2022b. Multiple brace root phenotypes promote anchorage and limit root lodging in maize. Plant, Cell & Environment, 45, 1573–1583.

Ishfaq S, Ding Y, Liang X Y, Guo W. 2025. Advancing lodging resistance in maize: Integrating genetic, hormonal, and agronomic insights for sustainable crop productivity. Plant Stress, 15, 100777.

Ivanchenko M G, Muday G K, Dubrovsky J G. 2008. Ethylene-auxin interactions regulate lateral root initiation and emergence in Arabidopsis thaliana. The Plant Journal, 55, 335–347.

Jia Q M, Xu Y Y, Ali S, Sun L F, Ding R X, Ren X L, Zhang P, Jia Z K. 2018. Strategies of supplemental irrigation and modified planting densities to improve the root growth and lodging resistance of maize (Zea mays L.) under the ridge-furrow rainfall harvesting system. Field Crops Research, 224, 48–59.

Kamara A Y, Kling J G, Menkir A, Ibikunle O. 2003. Association of vertical root-pulling resistance with root lodging and grain yield in selected S1 maize lines derived from a tropical low-nitrogen population. Journal of Agronomy and Crop Science, 189, 129–135.

Kriticos D J, Reynaud P, Baker R H A, Eyre D. 2012. Estimating the global area of potential establishment for the western corn rootworm (Diabrotica virgiferavirgifera) under rain-fed and irrigated agriculture: The potential distribution of western corn rootworm. Bulletin OEPP/EPPO Bulletin, 42, 56–64.

Kong X Z, Xiong Y L, Song X Y, Wadey S, Yu S H, Rao J L, Lale A, Lombardi M, Fusi R, Bhosale R, Huang G Q. 2023. Ethylene regulates auxin-mediated root gravitropic machinery and controls root angle in cereal crops. The Plant Cell, 35, 3560–3574.

Lakehal A, Bellini C. 2019. Control of adventitious root formation: Insights into synergistic and antagonistic hormonal interactions. Physiologia Plantarum, 165, 90–100.

Langan T D, Oplinger E S. 1987. Growth and yield of ethephon treated maize. Agronomy Journal, 79, 130–134.

Li S Y, Ma W, Peng J Y, Chen Z M. 2015. Study on yield loss of summer maize due to lodging at the big flare stage and grain filling stage. Scientia Agricultura Sinica, 19, 3952–3964. (in Chinese)

Liu B, He J X, Zeng F J, Lei J Q, Arndt S K. 2016. Life span and structure of ephemeral root modules of different functional groups from a desert system. New Phytologist, 211, 103–112.

Liu S Q, Song F B, Liu F L, Zhu X C, Xu H B. 2012. Effect of planting density on root lodging resistance and its relationship to nodal root growth characteristics in maize (Zea mays L.). Journal of Agricultural Science, 4, 182.

Liu S S, Ji X D, Zhang X. 2022. Effects of soil properties and tree species on root–soil anchorage characteristics. Sustainability, 14, 7770.

Lynch J P. 2019. Root phenotypes for improved nutrient capture: An underexploited opportunity for global agriculture. New Phytologist, 223, 548–564.

Lynch J P. 2013. Steep, cheap and deep: An ideotype to optimize water and N acquisition by maize root systems. Annals of Botany, 112, 347–357.

Lynch J P, Brown K M. 2012. New roots for agriculture: Exploiting the root phenome. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 367, 1598–1604.

Lynch J P, Chimungu J G, Brown K M. 2014. Root anatomical phenes associated with water acquisition from drying soil: Targets for crop improvement. Journal of Experimental Botany, 65, 6155–6166.

Mi G H, Chen F J, Wu Q P, Lai N W, Yuan L X, Zhang F S. 2010. Ideotype root architecture for efficient nitrogen acquisition by maize in intensive cropping systems. Science China Life Sciences, 53, 1369–1373.

Novacek M J, Mason S C, Galusha T D, Yaseen M. 2013. Twin rows minimally impact irrigated maize yield, morphology, and lodging. Agronomy Journal, 105, 268–276.

Qi X L, Cui K H, Liu L, Huang H Y, Xu Z Y, Wang Z M, Xiong D L, Huang J L, Peng S B. 2025. Exogenous ethephon application promotes nitrogen accumulation by modifying root characters in rice seedlings. The Crop Journal, 13, 1642–1647.

Qin X L, Feng F, Li Y J, Xu S T, Siddique K H M, Liao Y C. 2016. Maize yield improvements in China: Past trends and future directions. Plant Breeding, 135, 166–176.

Ren H, Liu Z, Wang X B, Zhou W B, Zhou B Y, Zhao M, Li C F. 2025. Long-term excessive nitrogen application decreases spring maize nitrogen use efficiency via suppressing root physiological characteristics. Journal of Integrative Agriculture, 24, 4195–4210.

Sanchez G M, Nejadhashemi A P, Zhang Z, Marquart-Pyatt S, Habron G, Shortridge A. 2015. Linking watershed-scale stream health and socioeconomic indicators with spatial clustering and structural equation modeling. Environmental Modelling & Software, 70, 113–127.

Sha X Q, Guan H H, Zhou Y Q, Su E H, Guo J, Li Y X, Zhang D F, Liu X Y, He G H, Li Y, Wang T Y, Zou H W, Li C H. 2023. Genetic dissection of crown root traits and their relationships with aboveground agronomic traits in maize. Journal of Integrative Agriculture, 22, 3394–3407.

Shao H, Shi D F, Shi W J, Ban X B, Chen Y C, Ren W, Chen F J, Mi G H. 2019. Genotypic difference in the plasticity of root system architecture of field-grown maize in response to plant density. Plant and Soil, 439, 201–217.

Shekoofa A, Emam Y. 2008. Effect of water stress, plant density and plant growth regulator (ethephon) on growth and yield of maize (Zea mays L.). Journal of Agronomy, 7, 41–48.

Shi J R, Drummond B J, Habben J E, Brugire N, Weers B P, Hakimi S M, Lafitte H R, Schussler J R, Mo H, Beatty M, Zastrow-Hayes G, O’Neill D. 2019. Ectopic expression of ARGOS8 reveals a role for ethylene in root-lodging resistance in maize. The Plant Journal, 97, 378–390.

Shi J R, Habben J E, Archibald R L, Drummond B J, Chamberlin M A, Williams R W, Lafitte H R, Weers B P. 2015. Overexpression of ARGOS genes modifies plant sensitivity to ethylene, leading to improved drought tolerance in both Arabidopsis and maize. Plant Physiology, 169, 266–282.

Smith S, De Smet I. 2012. Root system architecture: Insights from Arabidopsis and cereal crops. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 367, 1441–1452.

Sparks E E. 2023. Maize plants and the brace roots that support them. New Phytologist, 237, 48–52.

Stamp P, Kiel C. 1992. Root morphology of maize and its relationship to root lodging. Journal of Agronomy and Crop Science, 168, 113–118.

Steffens B, Rasmussen A. 2016. The physiology of adventitious roots. Plant Physiology, 170, 603–617.

Stepanova A N, Alonso J M. 2019. From ethylene-auxin interactions to auxin biosynthesis and signal integration. The Plant Cell, 31, 1393–1394.

Sun N, Chen X F, Zhao H X, Meng X M, Bian S F. 2022. Effects of plant growth regulators and nitrogen management on root lodging resistance and grain yield under high-density maize crops. Agronomy (Basel), 12, 2892.

Sun S X, Dai J Y. 1989. Effect of nitrogen, phosphate and potash fertilizers on lodging and yield in maize. Scientia Agricultura Sinica, 22, 28–33. (in Chinese)

Tilman D, Balzer C, Hill J, Befort B L. 2011. Global food demand and the sustainable intensification of agriculture. Proceedings of the National Academy of Sciences of the United States of America, 108, 20260–20264.

Torres-Martínez H H, Rodríguez-Alonso G, Shishkova S, Dubrovsky J G. 2019. Lateral root primordium morphogenesis in angiosperms. Frontiers in Plant Science, 10, 206.

Trachsel S, Kaeppler S M, Brown K M, Lynch J P. 2013. Maize root growth angles become steeper under low N conditions. Field Crops Research, 140, 18–31.

Wang X L, Liu F, Zhao N, Du X, Yin P J, Li T L, Lan T Q, Feng D J, Kong F L, Yuan J C. 2025. Optimizing sowing dates increase solar radiation to mitigate maize lodging and yield variability: A five-year field study. Journal of Integrative Agriculture, 24, 4573–4587.

Wang Z, Liu Z, Liu R X, Zhou B Y, Wang X B, Ding Z S, Zhang R H, Li C F, Zhao M. 2025. Ethephon combined with synergist promoted grain formation by improving photosynthetic performance and floret development of maize. Field Crops Research, 334, 110140.

Wei H Y, Hu L, Zhu Y, Xu D, Zheng L M, Chen Z F, Hu Y J, Cui P Y, Guo B W, Dai Q G, Zhang H C. 2018. Different characteristics of nutrient absorption and utilization between inbred japonica super rice and inter-sub-specific hybrid super rice. Field Crops Research, 218, 88–96.

White P J, George T S, Dupuy L X, Karley A J, Valentine T A, Wiesel L, Wishart J. 2013. Root traits for infertile soils. Frontiers in Plant Science, 4, 193.

Wiersma J J, Dai J, Durgan B R. 2011. Optimum timing and rate of trinexapac-ethyl to reduce lodging in spring wheat. Agronomy Journal, 103, 864–870.

Xu C L, Gao Y B, Tian B J, Ren J H, Meng Q F, Wang P. 2017. Effects of EDAH, a novel plant growth regulator, on mechanical strength, stalk vascular bundles and grain yield of summer maize at high densities. Field Crops Research, 200, 71–79.

Xue J, Xie R Z, Zhang W F, Wang K R, Hou P, Ming B, Gou L, Li S K. 2017. Research progress on reduced lodging of high-yield and-density maize. Journal of Integrative Agriculture, 16, 2717–2725.

Xue J, Gao S, Fan Y H, Li L L, Ming B, Wang K R, Xie R Z, Hou P, Li S K. 2020. Traits of plant morphology, stalk mechanical strength, and biomass accumulation in the selection of lodging-resistance maize cultivars. European Journal of Agronomy, 117, 126073.

Ye D L, Zhang Y S, Al-Kaisi M M, Duan L S, Zhang M C, Li Z H. 2016. Ethephon improved stalk strength associated with summer maize adaptations to environments differing in nitrogen availability in the North China Plain. The Journal of Agricultural Science, 154, 960–977.

Yin S Y, Li P C, Xu Y, Xue L, Hao D R, Liu J, Yang T T, Yang Z F, Xu C W. 2018. Logistic model-based genetic analysis for kernel filling in a maize RIL population. Euphytica, 214, 86.

York L M, Nord E A, Lynch J P. 2013. Integration of root phenes for soil resource acquisition. Frontiers in Plant Science, 4, 355.

Zhang H X, Ali S, Chen Y, Yang L Y, Pang G, Assal M E, Shaik M R. 2025. Irrigation management practices with novel plant growth regulators improve root growth, root lodging resistance and maize productivity under semi-arid regions. Field Crops Research, 333, 110097.

Zhang P, Gu S C, Wang Y Y, Yang R M, Yan Y, Zhang S, Sheng D C, Cui T, Huang S B, Wang P. 2021. Morphological and mechanical variables associated with lodging in maize (Zea mays L.). Field Crops Research, 269, 108178.

Zhang P, Wang Y Y, Sheng D C, Zhang S, Gu S C, Yan Y, Zhao F C, Wang P, Huang S B. 2023. Optimizing root system architecture to improve root anchorage strength and nitrogen absorption capacity under high plant density in maize. Field Crops Research, 303, 109109.

Zhang P, Yan Y, Gu S C, Wang Y Y, Xu C L, Sheng D C, Li Y B, Wang P, Huang S B. 2022. Lodging resistance in maize: A function of root–shoot interactions. European Journal of Agronomy, 132, 126393.

Zhang Q, Zhang L Z, Evers J, Werf W, Zhang W Q, Duan L S. 2014. Maize yield and quality in response to plant density and application of a novel plant growth regulator. Field Crops Research, 164, 82–89.

Zhang W Q, Yu C X, Zhang K, Zhou Y Y, Tan W M, Zhang L Z, Li Z H, Duan L S. 2017. Plant growth regulator and its interactions with environment and genotype affect maize optimal plant density and yield. European Journal of Agronomy, 91, 34–43.

Zhang Y S, Wang Y B, Ye D L, Wang W, Qiu X M, Duan L S, Li Z H, Zhang M C. 2019. Ethephon improved stalk strength of maize (Zea mays L.) mainly through altering internode morphological traits to modulate mechanical properties under field conditions. Agronomy (Basel), 9, 186.

Zhang Y S, Wang Y B, Ye D L, Xing J P, Duan L S, Li Z H, Zhang M C. 2020. Ethephon-regulated maize internode elongation associated with modulating auxin and gibberellin signal to alter cell wall biosynthesis and modification. Plant Science, 290, 110196.

Zheng Z G, Wang B B, Zhuo C Y, Xie Y R, Zhang X M, Liu Y J, Zhang G S, Ding H, Zhao B B, Tian M Q, Xu M Y, Kong D X, Shen R X, Liu Q, Wu G X, Huang J F, Wang H Y. 2023. Local auxin biosynthesis regulates brace root angle and lodging resistance in maize. New Phytologist, 238, 142–154.

Zhu Y P, Wang S, Li Y H, Wei D, Luo N, Wang P, Meng Q F. 2023. Enhancing maize yield through understanding the novel shoot-root interaction in morphology among hybrids with dense planting. Field Crops Research, 302, 109107.