Acorn size is more important than nursery fertilization for outplanting performance of Quercus variabilis container seedlings

  • Wenhui Shi
  • Pedro Villar-Salvador
  • Guolei LiEmail author
  • Xiaoxu Jiang
Research Paper


Key message

Small acorns are usually discarded for seedling cultivation because they reduce plant quality. This, however, can potentially reduce genetic diversity of plantations. The use of small acorns will result in the production of a higher proportion of small seedlings containing low nutrient levels and having poor outplanting performance in oak container seedlings. Nursery fertilization partially offsets the negative effect of small acorns on seedling attributes in the nursery but not on outplanting performance.


Small acorns result in low-quality seedlings and so are usually discarded in artificial regeneration programs of oak species. This can potentially reduce genetic diversity of plantations. Nursery fertilization may compensate for the low quality of small-acorn seedlings.


To assess whether nursery fertilization interacts with Quercus variabilis acorn size to determine seedling morphology and nutrition in the nursery and outplanting performance.


Acorns of three size classes were used to cultivate seedlings with or without fertilization. Seedling emergence, nursery morphology and nutrient status, and outplanting survival and growth were measured.


Small acorns represented 41% of the seed batch. Most acorn size variation occurred within trees rather than among trees. Smaller acorns were associated with lower emergence and resulted in smaller seedlings that had lower nutrient content levels. Nursery fertilization slightly increased seedling growth for all acorn sizes; it also strongly increased nutrient content, especially in small-acorn seedlings. Two years after planting, survival of small-acorn seedlings was 32% lower than the survival of medium- and large-acorn plants. Fertilization did not affect survival, but it did increase size, especially of small-acorn seedlings, though they did not achieve the growth of large-acorn seedlings.


Nursery fertilization increases growth and nutrient status, but not outplanting performance, of small-acorn seedlings.


Acorn size Nursery fertilization Nutrient concentration Outplanting performance Seedling growth Survival 



We thank the managers and workers at the Jiufeng Mountain greenhouse for their valuable help and support.


This work was funded by the the National Natural Science Foundation of China (31670638), Fundamental Research Funds for Central Universities (2017PT02), Special Fund for Beijing Common Construction Project, Project CGL 2014-53308-P of the Spanish government, and the REMEDINAL 3 network (S2013/MAE-2719) of the CAM.

Data availability

The datasets generated and analyzed in the current study are available in the Mendeley repository (Shi et al. 2018b). Datasets are not peer-reviewed. Shi et al. (2018b) Data from: Acorn size is more important than nursery fertilization for outplanting performance of Quercus variabilis container seedlings. V2. Mendeley Digital Repository. [dataset].

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflicts of interest.


  1. Bonfil C (1998) The effects of seed size, cotyledon reserves, and herbivory on seedling survival and growth in Quercus rugosa and Q. Laurina (Fagaceae). Am J Bot 85:79–87CrossRefGoogle Scholar
  2. Bonner FT (2008) Quercus. In: Bonner FT, Karrfalt RP (eds) The woody plant seed manual, USDA Forest Service, agriculture handbook, vol 727, pp 928–938Google Scholar
  3. Burgarella C, Navascues M, Soto Á et al (2007) Narrow genetic base in forest restoration with holm oak (Quercus ilex L.) in Sicily. Ann For Sci 64:757–763CrossRefGoogle Scholar
  4. Carlo TA, Yang S (2011) Network models of frugivory and seed dispersal: challenges and opportunities. Acta Oecol 37:619–624CrossRefGoogle Scholar
  5. Climent J, Chambel MR, Pardos M, Lario F, Villar-Salvador P (2011) Biomass allocation and foliage heteroblasty in hard pine species respond differentially to reduction in rooting volume. Eur J For Res 130:841–850CrossRefGoogle Scholar
  6. Connor KF, Sowa S (2003) Effects of desiccation on the physiology and biochemistry of Quercus alba acorns. Tree Physiol 23:1147–1152CrossRefGoogle Scholar
  7. Cuesta B, Villar-Salvador P, Puértolas J, Rey Benayas JM, Michalet R (2010) Facilitation of Quercus ilex in Mediterranean shrubland is explained by both direct and indirect interactions mediated by herbs. J Ecol 98:687–696CrossRefGoogle Scholar
  8. Flint SD, Palmblad IG (1978) Germination dimorphism and developmental flexibility in the ruderal weed Heterotheca grandiflora. Oecologia 36:33–43CrossRefGoogle Scholar
  9. Foster SA, Janson CH (1985) The relationship between seed size and establishment conditions in tropical woody plants. Ecology 66:773–780CrossRefGoogle Scholar
  10. Ganatsas P, Tsakaldimi M (2013) A comparative study of desiccation responses of seeds of three drought-resistant Mediterranean oaks. For Ecol Manag 305:189–194CrossRefGoogle Scholar
  11. Giertych MJ, Suszka J (2011) Consequences of cutting off distal ends of cotyledons of Quercus robur acorns before sowing. Ann For Sci 68:433–442CrossRefGoogle Scholar
  12. Gil-Pelegrín E, Peguero-Pina JJ, Sancho-Knapik D (2017) Oaks and people: a long journey together. In: Gil-Pelegrín E, Peguero-Pina JJ, Sancho-Knapik D (eds) Oaks physiological ecology. Exploring the functional diversity of Genus Quercus L. Springer International Publishing, Cham, pp 1–11Google Scholar
  13. González-Rodríguez V, Villar R, Navarro-Cerrillo RM (2011a) Maternal influences on seed mass effect and initial seedling growth in four Quercus species. Acta Oecol 37:1–9CrossRefGoogle Scholar
  14. González-Rodríguez V, Navarro-Cerrillo RM, Villar R (2011b) Artificial regeneration with Quercus ilex L. and Quercus suber L. by direct seeding and planting in southern Spain. Ann For Sci 68:637–646CrossRefGoogle Scholar
  15. Grossnickle SC (2012) Why seedlings survive: influence of plant attributes. New For 43:711–738CrossRefGoogle Scholar
  16. Grossnickle SC, Ivetić V (2017) Direct seeding in reforestation – a field performance review. Reforesta 4:94–142CrossRefGoogle Scholar
  17. Grossnickle SC, MacDonald JE (2018) Why seedlings grow: influence of plant attributes. New For 49:1–34CrossRefGoogle Scholar
  18. Johnson PS, Shifley SR, Rogers R (2009) The ecology and silviculture of oaks, 2nd edn. MPG Books Group, Bodmin, UKCrossRefGoogle Scholar
  19. Ke G, Werger MJA (1999) Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size. Acta Oecol 20:579–586CrossRefGoogle Scholar
  20. Knoot TG, Schulte LA, Rickenbach M (2010) Oak conservation and restoration on private forestlands: negotiating a social-ecological landscape. Environ Manag 45:155–164CrossRefGoogle Scholar
  21. Kormanik PP, Sung SS, Kormanik TL, Schlarbaum SE, Zarnoch SJ (1998) Effect of acorn size on development of northern red oak 1-0 seedlings. Can J For Res 28:1805–1813CrossRefGoogle Scholar
  22. Leishman MR, Westoby M, Jurado E (1995) Correlates of seed size variation: a comparison among five temperate floras. J Ecol 83:517–529CrossRefGoogle Scholar
  23. Leishman MR, Wright IJ, Moles AT, Westoby M (2000) The evolutionary ecology of seed size. In: Fenner M (ed) Seeds: the ecology of regeneration in plant communities, 2nd edn. CAB International, Wallingford, pp 31–57CrossRefGoogle Scholar
  24. Li G, Zhu Y, Liu Y, Wang J, Liu J, Dumroese RK (2014) Combined effects of pre-hardening and fall fertilization on nitrogen translocation and storage in Quercus variabilis seedlings. Eur J For Res 133:983–992CrossRefGoogle Scholar
  25. Li G, Liu Y, Zhu Y et al (2012) A review on the abroad studies of techniques in regulating quality of container seedling. Sci Silvae Sin 48:135–142 (In Chinese with English summary)Google Scholar
  26. Liu Y, Liu G, Li Q, et al (2012) Influence of pericarp, cotyledon and inhibitory substances on sharp tooth oak (Quercus aliena var. acuteserrata) germination. PLoS one.
  27. Mendez M (1997) Sources of variation in seed mass in Arum italicum. Int J Plant Sci 158:298–305CrossRefGoogle Scholar
  28. Moles A, Leishman M (2008) The seedling as part of a plant’s life history strategy. In: Allessio LM, Parker VT, Simpson RL (eds) Seedling ecology and evolution. Cambridge University Press, New York, pp 215–235Google Scholar
  29. Navarro FB, Jiménez MN, Ripoll MÁ, Fernández-Ondoño E, Gallego E, de Simón E (2006) Direct sowing of holm oak acorns: effects of acorn size and soil treatment. Ann For Sci 63:961–967CrossRefGoogle Scholar
  30. Oliet JA, Tejada M, Salifu KF, Collazos A, Jacobs DF (2009) Performance and nutrient dynamics of holm oak (Quercus ilex L.) seedlings in relation to nursery nutrient loading and post-transplant fertility. Eur J For Res 128:253–263CrossRefGoogle Scholar
  31. Pemán J, Chirino E, Espelta JM, et al (2017) Physiological keys for natural and artificial regeneration of oaks. In: Gil-Pelegrín E, Peguero-Pina J, Sancho-Knapik D (eds) Oaks physiological ecology. Exploring the functional diversity of Genus Quercus L. Springer, Cham, pp 453–511Google Scholar
  32. Pesendorfer MB (2014) The effect of seed size variation in Quercus pacifica on seedling establishment and growth. Gen Tech Rep PSW-GTR-251 407–412Google Scholar
  33. Pizo MA, Von Allmen C, Morellato LPC (2006) Seed size variation in the palm Euterpe edulis and the effects of seed predators on germination and seedling survival. Acta Oecol 29:311–315CrossRefGoogle Scholar
  34. Quero JL, Villar R, Marañón T et al (2007) Seed-mass effects in four Mediterranean Quercus species (Fagaceae) growing in contrasting light environments. Am J Bot 94:1795–1803CrossRefGoogle Scholar
  35. Ramírez-Valiente JA, Valladares F, Gil L, Aranda I (2009) Population differences in juvenile survival under increasing drought are mediated by seed size in cork oak (Quercus suber L.). For Ecol Manag 257:1676–1683CrossRefGoogle Scholar
  36. Rice KJ, Gordon DR, Hardison JL, Welker JM (1993) Phenotypic variation in seedlings of a “keystone” tree species (Quercus douglasii): the interactive effects of acorn source and competitive environment. Oecologia 96:537–547CrossRefGoogle Scholar
  37. Sage RD, Koenig WD, McLaughlin BC (2011) Fitness consequences of seed size in the valley oak Quercus lobata Née (Fagaceae). Ann For Sci 68:477–484CrossRefGoogle Scholar
  38. Salifu KF, Timmer VR (2003) Optimizing nitrogen loading of Picea mariana seedlings during nursery culture. Can J For Res 33:1287–1294CrossRefGoogle Scholar
  39. Salifu KF, Jacobs DF, Birge ZKD (2009) Nursery nitrogen loading improves field performance of bareroot oak seedlings planted on abandoned mine lands. Restor Ecol 17:339–349CrossRefGoogle Scholar
  40. Sánchez-Montes de Oca EJ, Badano EI, Silva-Alvarado LE et al (2018) Acorn weight as determinant of germination in red and white oaks: evidences from a common-garden greenhouse experiment. Ann For Sci 75:1–12CrossRefGoogle Scholar
  41. Shi W, Bloomberg M, Li G, Su S, Jia L (2017) Combined effects of cotyledon excision and nursery fertilization on root growth, nutrient status and outplanting performance of Quercus variabilis container seedlings. PLoS One 12:e0177002CrossRefGoogle Scholar
  42. Shi W, Villar-Salvador P, Jacobs DF, Li G, Jiang X (2018a) Simulated predation of Quercus variabilis acorns impairs nutrient remobilization and seedling performance irrespective of soil fertility. Plant Soil 423:295–306Google Scholar
  43. Shi W, Villar-Salvador P, Li G, et al (2018b) Data from: acorn size is more important than nursery fertilization for outplanting performance of Quercus variabilis container seedlings. V2. Mendeley digital repository. [dataset].
  44. Sung SS, Kormanik PP, Cook CD, et al (2006) Effect of acorn moisture content at sowing on germination and seedling growth of white oak and northern red oak. In: Gen. Tech. Rep. SRS-92. Department of Agriculture, Forest Service, Southern Research Station, Asheville, NC: U.S., pp 241–246Google Scholar
  45. Tecklin J, McCreary DD (1991) Acorn size as a factor in early seedling growth. In: Standiford RB (ed) Proceedings of the symposium on oak woodlands and hardwood rangeland management. Gen. Tech. Rep. PSW-126. CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, pp 48–53Google Scholar
  46. Tripathi RS, Khan ML (1990) Effects of seed weight and microsite characteristics on germination and seedling fitness in two species of Quercus in a subtropical wet hill forest. Oikos 57:289–296CrossRefGoogle Scholar
  47. Tungate KD, Burton MG, Susko DJ, Sermons SM, Rufty TW (2006) Altered weed reproduction and maternal effects under low-nitrogen fertility. Weed Sci 54:847–853. CrossRefGoogle Scholar
  48. Venable DL, Brown JS (1988) The selective interactions of dispersal, dormancy, and seed size as adaptations for reducing risk in variable environments. Am Nat 131:360–384CrossRefGoogle Scholar
  49. Villar-Salvador P, Planelles R, Enríquez E, Rubira JP (2004) Nursery cultivation regimes, plant functional attributes, and field performance relationships in the Mediterranean oak Quercus ilex L. For Ecol Manag 196:257–266CrossRefGoogle Scholar
  50. Villar-Salvador P, Heredia N, Millard P (2010) Remobilization of acorn nitrogen for seedling growth in holm oak (Quercus ilex), cultivated with contrasting nutrient availability. Tree Physiol 30:257–263CrossRefGoogle Scholar
  51. Villar-Salvador P, Puértolas J, Cuesta B, Peñuelas JL, Uscola M, Heredia-Guerrero N, Rey Benayas JM (2012) Increase in size and nitrogen concentration enhances seedling survival in Mediterranean plantations. Insights from an ecophysiological conceptual model of plant survival. New For 43:755–770CrossRefGoogle Scholar
  52. Villar-Salvador P, Peñuelas JL, Nicolás-Peragón JL, Benito LF, Domínguez-Lerena S (2013) Is nitrogen fertilization in the nursery a suitable tool for enhancing the performance of Mediterranean oak plantations? New For 44:733–751CrossRefGoogle Scholar
  53. Wagstaff K, Cardie C, Rogers S, Schroedl S (2001) Constrained K-means clustering with background knowledge. Int Conf Mach Learn 577–584Google Scholar
  54. Wang X (2013) Indicating function of Shanxi Zhongtiaoshan mountain oak forests to forest restoration and reconstruction in North China. For Econ 8:74–76Google Scholar
  55. Westoby M, Jurado E, Leishman M (1992) Comparative evolutionary ecology of seed size. Trends Ecol Evol 7:368–372CrossRefGoogle Scholar
  56. Westoby M, Leishman M, Lord J (1996) Comparative ecology of seed size and dispersal. Philos Trans R Soc B Biol Sci 351:1309–1318CrossRefGoogle Scholar
  57. Willis SG, Hulme PE (2004) Environmental severity and variation in the reproductive traits of Impatiens glandulifera. Funct Ecol 18:887–898CrossRefGoogle Scholar
  58. Wulff RD (1986) Seed size variation in Desmodium paniculatum: I. Factors affecting seed size. Ecology 74:87–97CrossRefGoogle Scholar
  59. Zhang W, Lu Z, Li J, Liu G (2002) A comparative study on spatial distribution pattern and its dynamics of Quercus variabilis populations among different forest areas in Shanxi province, China. Acta Bot Boreali-occidentalia Sin 22:476–483 (In Chinese with English summary)Google Scholar

Copyright information

© INRA and Springer-Verlag France SAS, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Key Laboratory for Silviculture and Conservation, Ministry of EducationBeijing Forestry UniversityBeijingChina
  2. 2.State Key Laboratory of Subtropical SilvicultureZhejiang A&F UniversityHangzhouChina
  3. 3.Forest Ecology and Restoration Group, Departamento de Ciencias de la VidaUniversidad de AlcaláAlcalá de HenaresSpain
  4. 4.Beijing Laboratory of Urban and Rural Ecological EnvironmentBeijing Municipal Education CommissionBeijingChina

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