Plant and Soil

, Volume 430, Issue 1–2, pp 395–411 | Cite as

Root growth and root system architecture of field-grown maize in response to high planting density

  • Hui Shao
  • Tingting Xia
  • Dali Wu
  • Fanjun Chen
  • Guohua MiEmail author
Regular Article



This paper aims to investigate the adaptation of maize root system architecture (RSA) in response to increasing planting densities.


A three-year field study was conducted with three planting densities (40,000, 70,000, and 90,000 plants per ha, which are abbreviated as D40000, D70000 and D90000, respectively). The dynamic change of root morphological traits and the 3-dimensional RSA were quantified.


The grain yield per ha increased with increasing plant density from D40000 to D70000, and then decreased at D90000. Compared to D70000, high planting density of D90000 did not changed the total root biomass per ha but increased shoot biomass per ha by 4 to 8% in two of the three experimental years. Grain yield per plant and plant NPK concentration decreased with increasing planting density. Total accumulation of P and K per ha also decreased at D90000 compared to D70000. Root to shoot ratio was reduced at high planting density beginning 50 days after emergence. Compared to the control (D70000), total root length (TRL) per plant was reduced by 18 to 30% at D90000 and increased by 43 to 56% at D40000, root biomass per plant was reduced by 23 to 34% at D90000 and increased by 66 to 75% at D40000. High plant density reduced the number of nodal roots, lateral root density (LRD) and the average lateral root (LR) length, but with less effect on the length of axial roots. The RSA is characteristic of “intra-row contraction and inter-row extension”. Vertically, root growth in top soil layer (0- to 36- cm) was enhanced under supra-optimal plant density, but had a negligible effect in deep soil layers (36- to 60- cm).


To adapt to the limited photosynthesis capacity in the roots under high planting density, maize plants tend to reduce nodal root number and inhibit lateral root growth. They maintain nodal root length to explore a larger soil space, and adjust root growth in the intra-row and inter-row direction to avoid root-to-root competition.


Inter-row Intra-row Root system architecture Competition Plant density Maize 



40,000 plants per ha


70,000 plants per ha


90,000 plants per ha


total root length


lateral roots


lateral root density


specific root length


root length density



This work was financially supported by National Basic Research Program (973 Program) of China (2015CB150402) and State key research program (2016YFD0300304).

Supplementary material

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  1. Asseng S, Ritchie J, Smucker A, Robertson M (1998) Root growth and water uptake during water deficit and recovering in wheat. Plant Soil 201(2):265–273CrossRefGoogle Scholar
  2. Barber S et al (1988) Effects of soil temperature and water on maize root growth. Plant Soil 111(2):267–269CrossRefGoogle Scholar
  3. Biedrzycki ML, Jilany TA, Dudley SA, Bais HP (2010) Root exudates mediate kin recognition in plants. Communicative & integrative biology 3(1):28–35CrossRefGoogle Scholar
  4. Biernaskie JM (2011) Evidence for competition and cooperation among climbing plants. Proc R Soc Lond B Biol Sci 278(1714):1989–1996CrossRefGoogle Scholar
  5. Böhm W (1979) Methods of studying root systems. Ecological studies, 33. Springer, Berlin, pp 20–25Google Scholar
  6. Chen BJ, During HJ, Anten NP (2012a) Detect thy neighbor: identity recognition at the root level in plants. Plant Sci 195:157–167CrossRefPubMedGoogle Scholar
  7. Chen YL et al (2012b) Root growth and its response to increasing planting density in differentmaize hybrids. Plant Nutrition and Fertilizer Science 18(1):52–59 in ChineseGoogle Scholar
  8. Chen YL et al (2014) Characterization of the plant traits contributed to high grain yield and high grain nitrogen concentration in maize. Field Crop Res 159:1–9CrossRefGoogle Scholar
  9. Crepy MA, Casal JJ (2015) Photoreceptor-mediated kin recognition in plants. New Phytol 205(1):329–338CrossRefPubMedGoogle Scholar
  10. Cui Z, Chen X, Miao Y, Zhang F, Sun Q, Schroder J, Xu J (2008) On-farm evaluation of the improved soil N min–based nitrogen management for summer maize in North China plain. Agron J 100(3):517–525CrossRefGoogle Scholar
  11. D’Andrea K et al (2008) Kernel number determination differs among maize hybrids in response to nitrogen. Field Crop Res 105(3):228–239CrossRefGoogle Scholar
  12. Demotes-Mainard S, Pellerin S (1992) Effect of mutual shading on the emergence of nodal roots and the root/shoot ratio of maize. Plant Soil 147(1):87–93CrossRefGoogle Scholar
  13. Donald CT (1968) The breeding of crop ideotypes. Euphytica 17(3):385–403CrossRefGoogle Scholar
  14. Dudley SA, File AL (2007) Kin recognition in an annual plant. Biol Lett 3(4):435–438CrossRefPubMedPubMedCentralGoogle Scholar
  15. Dunbabin V et al (2003) Is there an optimal root architecture for nitrate capture in leaching environments? Plant Cell Environ 26(6):835–844CrossRefPubMedGoogle Scholar
  16. Eissenstat DM (1992) Costs and benefits of constructing roots of small diameter. J Plant Nutr 15(6–7):763–782CrossRefGoogle Scholar
  17. Ennos A et al (1993) The anchorage mechanics of maize, Zea mays. J Exp Bot 44(1):147–153CrossRefGoogle Scholar
  18. Feldman L (1994) The maize root. In: The maize handbook. Springer, New York, pp 29–37CrossRefGoogle Scholar
  19. File, A. L., Murphy, G. P., & Dudley, S. A. (2012). Fitness consequences of plants growing with siblings: reconciling kin selection, niche partitioning and competitive ability. Paper presented at the Proc. R. Soc. BGoogle Scholar
  20. Girardin P (1992) Leaf azimuth in maize canopies. Eur J Agron 1(2):91–97CrossRefGoogle Scholar
  21. Girardin P, Tollenaar M (1994) Effects of intraspecific interference on maize leaf azimuth. Crop Sci 34(1):151–155CrossRefGoogle Scholar
  22. Goodman AM, Ennos AR (1997) The responses of field-grown sunflower and maize to mechanical support. Ann Bot 79(6):703–711CrossRefGoogle Scholar
  23. Granato TC, Raper CD (1989) Proliferation of maize (Zea mays L.) roots in response to localized supply of nitrate. J Exp Bot 40(2):263–275CrossRefPubMedGoogle Scholar
  24. Grzesiak S et al (1999) The impact of limited soil moisture and waterlogging stress conditions on morphological and anatomical root traits in maize (Zea mays L.) hybrids of different drought tolerance. Acta Physiol Plant 21(3):305–315CrossRefGoogle Scholar
  25. Guingo E, Hébert Y (1997) Relationship between mechanical resistance of the maize root system and root morphology, and their genotypic and environmental variation. Maydica 42(3):265–274Google Scholar
  26. Hammer GL et al (2009) Can changes in canopy and/or root system architecture explain historical maize yield trends in the U.S. Corn Belt? Crop Sci 49(1):299–312CrossRefGoogle Scholar
  27. Hebert Y, et al (1992) Root lodging resistance in forage maize. Genetic variability of root system and aerial part. Maydica (Italy)Google Scholar
  28. Hebert Y et al (2001) The response of root/shoot partitioning and root morphology to light reduction in maize genotypes. Crop Sci 41(2):363–371CrossRefGoogle Scholar
  29. Hecht VL et al (2016) Sowing density: a neglected factor fundamentally affecting root distribution and biomass allocation of field grown spring barley (Hordeum vulgare L.). Front Plant Sci 7:944. CrossRefPubMedPubMedCentralGoogle Scholar
  30. Ho MD et al (2005) Root architectural tradeoffs for water and phosphorus acquisition. Funct Plant Biol 32(8):737–748CrossRefGoogle Scholar
  31. Hochholdinger F (2009) The maize root system: morphology, anatomy, and genetics. Handbook of maize: its biology. Springer, New York, pp 145–160CrossRefGoogle Scholar
  32. Hochholdinger F (2016) Untapping root system architecture for crop improvement [J]. J Exp Bot 67(15):4431CrossRefPubMedPubMedCentralGoogle Scholar
  33. Hochholdinger F et al (2004) From weeds to crops: genetic analysis of root development in cereals. Trends Plant Sci 9(1):42–48CrossRefPubMedGoogle Scholar
  34. Hoppe D et al (1986) The nodal roots of Zea: their development in relation to structural features of the stem. Can J Bot 64(11):2524–2537CrossRefGoogle Scholar
  35. Jansen C et al (2005) Investigating a trade-off in root morphological responses to a heterogeneous nutrient supply and to flooding. Funct Ecol 19(6):952–960CrossRefGoogle Scholar
  36. Kuchenbuch R, Barber S (1987) Yearly variation of root distribution with depth in relation to nutrient uptake and corn yield 1. Communications in Soil Science & Plant Analysis 18(3):255–263CrossRefGoogle Scholar
  37. Kumar B et al (2012) Genotypic variation for root architecture traits in seedlings of maize (Zea mays L.) inbred lines. Plant Breed 131(4):465–478CrossRefGoogle Scholar
  38. Lambers H, Posthumus F (1980) The effect of light intensity and relative humidity on growth rate and root respiration of Plantago lanceolata and Zea mays. J Exp Bot 31(6):1621–1630CrossRefGoogle Scholar
  39. Li T et al (2014) Effects of mutual shading on the regulation of photosynthesis in field-grown sorghum. J Photochem Photobiol B Biol 137:31–38CrossRefGoogle Scholar
  40. Liu S et al (2012) Effect of planting density on root lodging resistance and its relationship to nodal root growth characteristics in maize (Zea mays L.). J Agric Sci 4(12):182Google Scholar
  41. Lynch J (1995) Root architecture and plant productivity. Plant Physiol 109(1):7–13CrossRefPubMedPubMedCentralGoogle Scholar
  42. Lynch J, Brown K (1998) Regulation of root architecture by phosphorus availability. Current Topics Plant Physiol 19:148–156Google Scholar
  43. Lynch JP, Brown KM (2001) Topsoil foraging - an architectural adaptation of plants to low phosphorus availability. Plant Soil 237(2):225–237CrossRefGoogle Scholar
  44. Lynch JP, Brown KM (2008) Root strategies for phosphorus acquisition. The ecophysiology of plant-phosphorus interactions. Springer, Netherlands, pp 83–116CrossRefGoogle Scholar
  45. Lynch JP, Brown KM (2012) New roots for agriculture: exploiting the root phenome. Philos Trans R Soc Lond 367(1595):1598–1604CrossRefGoogle Scholar
  46. Lynch JP, Ho MD (2005) Rhizoeconomics: carbon costs of phosphorus acquisition. Plant Soil 269(1):45–56CrossRefGoogle Scholar
  47. Lynch, J.P., Chimungu, J.G. and Brown, K.M. (2014) Root anatomical phenes associated with water acquisition from drying soil: targets for crop improvement. J Exp Bot 65(21):6155–6166Google Scholar
  48. Ma Q et al (2013) Localized application of NH4 +-N plus P at the seedling and later growth stages enhances nutrient uptake and maize yield by inducing lateral root proliferation. Plant Soil 372(1–2):65–80CrossRefGoogle Scholar
  49. Ma D et al (2014) Lodging-related stalk characteristics of maize varieties in China since the 1950s. Crop Sci 54(6):2805–2814CrossRefGoogle Scholar
  50. Maddonni GA et al (1998) Grain yield components in maize. II. Postsilking growth and kernel weight. Field Crop Res 56(3):257–264CrossRefGoogle Scholar
  51. Maddonni G et al (2001) Plant population density, row spacing and hybrid effects on maize canopy architecture and light attenuation. Field Crop Res 71(3):183–193CrossRefGoogle Scholar
  52. Maddonni GA, Otegui ME, Andrieu B, Chelle M, Casal JJ (2002) Maize leaves turn away from neighbors. Plant Physiol 130(3):1181–1189CrossRefPubMedPubMedCentralGoogle Scholar
  53. Marler TE (2013) Kin recognition alters root and whole plant growth of split-root Cycas edentata seedlings. HortScience 48(10):1266–1269Google Scholar
  54. Mi G et al (2010) Ideotype root architecture for efficient nitrogen acquisition by maize in intensive cropping systems. Sci China Life Sci 53(12):1369–1373CrossRefPubMedGoogle Scholar
  55. Mi G et al (2016) Ideotype root system architecture for maize to achieve high yield and resource use efficiency in intensive cropping systems. Adv Agron 139:73–97CrossRefGoogle Scholar
  56. Mollier A, Pellerin S (1999) Maize root system growth and development as influenced by phosphorus deficiency. J Exp Bot 50(333):487–497CrossRefGoogle Scholar
  57. Murphy GP, Dudley SA (2009) Kin recognition: competition and cooperation in Impatiens (Balsaminaceae). Am J Bot 96(11):1990–1996CrossRefPubMedGoogle Scholar
  58. Murphy GP, Van Acker R, Rajcan I, Swanton CJ (2017) Identity recognition in response to different levels of genetic relatedness in commercial soya bean. Royal Society open science 4(1):160879CrossRefPubMedPubMedCentralGoogle Scholar
  59. Nelson DW, Sommers LE (1973) Determination of total nitrogen in plant material. Agron J 65:109–112CrossRefGoogle Scholar
  60. Ning P et al (2012) Maize cob plus husks mimics the grain sink to stimulate nutrient uptake by roots. Field Crop Res 130(2):38–45CrossRefGoogle Scholar
  61. Ning P et al (2013) Post-silking accumulation and partitioning of dry matter, nitrogen, phosphorus and potassium in maize varieties differing in leaf longevity. Field Crop Res 144(144):19–27CrossRefGoogle Scholar
  62. Pellerin S, Demotes-Mainard S, Kutschera L (1992) Effect of competition for light between plants on the root shoot ratio and the number of adventitious roots of maize. In Proc 3:65–68Google Scholar
  63. Pinthus MJ (1967) Spread of the root system as indicator for evaluating lodging resistance of wheat. Crop Sci 7(2):107–110CrossRefGoogle Scholar
  64. Poorter H et al (2016) Pampered inside, pestered outside? Differences and similarities between plants growing in controlled conditions and in the field. New Phytol 212(4):838–855CrossRefPubMedGoogle Scholar
  65. Richardson AE et al (2011) Plant and microbial strategies to improve the phosphorus efficiency of agriculture. Plant Soil 349(1):121–156CrossRefGoogle Scholar
  66. Ryser P (2006) The mysterious root length. Plant Soil 286:1): 1–1): 6CrossRefGoogle Scholar
  67. Saengwilai P et al (2014) Low crown root number enhances nitrogen acquisition from low-nitrogen soils in maize. Plant Physiol 166(2):581–589CrossRefPubMedPubMedCentralGoogle Scholar
  68. Saini HS, Westgate ME (1999) Reproductive development in grain crops during drought. Adv Agron 68:59–96CrossRefGoogle Scholar
  69. Sangoi L (2001) Understanding plant density effects on maize growth and development: an important issue to maximize grain yield. Ciência Rural 31(1):159–168CrossRefGoogle Scholar
  70. Sanguineti M et al (1998) Root and shoot traits of maize inbred lines grown in the field and in hydroponic culture and their relationships with root lodging. Maydica 43:211–216Google Scholar
  71. Semchenko M, Saar S, Lepik A (2014) Plant root exudates mediate neighbour recognition and trigger complex behavioural changes. New Phytol 204(3):631–637CrossRefPubMedGoogle Scholar
  72. Singh V et al (2012) Genetic control of nodal root angle in sorghum and its implications on water extraction. Eur J Agron 42(5):3–10CrossRefGoogle Scholar
  73. Soil Survey Staff (1998) Keys to Soil Taxonomy. United States Department of Agriculture, Natural Resources Conservation Service, Washington, DC, USA, pp. 211Google Scholar
  74. Soon Y K, Kalra Y P. A. (1995) comparison of plant tissue digestion methods for nitrogen and phosphorus analyses [J]. Can J Soil Sci, 75(2): 243–245Google Scholar
  75. Stamp P, Kiel C (1992) Root morphology of maize and its relationship to root lodging. J Agron Crop Sci 168(2):113–118CrossRefGoogle Scholar
  76. Thorup-Kristensen K (2001) Are differences in root growth of nitrogen catch crops important for their ability to reduce soil nitrate-N content, and how can this be measured? Plant Soil 230(2):185–195CrossRefGoogle Scholar
  77. Tokatlidis I, Koutroubas S (2004) A review of maize hybrids’ dependence on high plant populations and its implications for crop yield stability. Field Crop Res 88(2):103–114CrossRefGoogle Scholar
  78. Trachsel S et al (2011) Shovelomics: high throughput phenotyping of maize (Zea mays L.) root architecture in the field. Plant Soil 341(1–2):75–87CrossRefGoogle Scholar
  79. Varney GT, Mccully ME (2010) The branch roots of Zea. II. Developmental loss of the apical meristem in field-grown roots. New Phytol 118(4):535–546CrossRefGoogle Scholar
  80. Wahl S, Ryser P (2000) Root tissue structure is linked to ecological strategies of grasses. New Phytol 148(3):459–471CrossRefGoogle Scholar
  81. Wang X et al (2014) Effect of strip subsoiling on population root spatial distribution of maize under different planting densities. Acta Agron Sin 40(12):2136–2148CrossRefGoogle Scholar
  82. Weiner J (2003) Ecology–the science of agriculture in the 21st century. J Agric Sci 141(3–4):371–377CrossRefGoogle Scholar
  83. Wiesler F, Horst W (1994) Root growth and nitrate utilization of maize cultivars under field conditions. Plant Soil 163(2):267–277CrossRefGoogle Scholar
  84. Xue J et al (2016a) Effects of light intensity within the canopy on maize lodging. Field Crop Res 188:133–141CrossRefGoogle Scholar
  85. Xue J et al (2016b) How high plant density of maize affects basal internode development and strength formation. Crop Sci 56(6):3295CrossRefGoogle Scholar
  86. York LM et al (2015) Evolution of US maize (Zea mays L.) root architectural and anatomical phenes over the past 100 years corresponds to increased tolerance of nitrogen stress. J Exp Bot 66(8):2347–2358CrossRefPubMedPubMedCentralGoogle Scholar
  87. Yu DW (2014) Study on the effect of brace root traits on lodging resistance and genetic research of maize brace root traits.[D]. North West Agriculture and Forestry University, Yangling (in Chinese)Google Scholar
  88. Zhang R et al (2015) Effect of subsoiling on root morphological and physiological characteristics of spring maize. transactions of the Chinese society of. Agric Eng 31(5):78–84Google Scholar
  89. Zhu L, Zhang D-Y (2013) Donald’s ideotype and growth redundancy: a pot experimental test using an old and a modern spring wheat cultivar. PLoS One 8(7):e70006CrossRefPubMedPubMedCentralGoogle Scholar
  90. Zhu J et al (2005) Topsoil foraging and phosphorus acquisition efficiency in maize (Zea mays). Funct Plant Biol 32(8):749–762CrossRefGoogle Scholar

Copyright information

© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  • Hui Shao
    • 1
  • Tingting Xia
    • 1
  • Dali Wu
    • 1
  • Fanjun Chen
    • 1
  • Guohua Mi
    • 1
    Email author
  1. 1.Center for Resources, Environment and Food Security, College of Resources and Environmental ScienceChina Agricultural UniversityBeijingPeople’s Republic of China

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