Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Growth responses to irradiance regime along an ecological gradient of Quercus castaneifolia seedlings of different provenance


Understory light is known as one of the most important environmental predictors of growth response of woody species. Hence, the primary objective of most forest management practices is optimizing that resource for understory seedlings. In this study, growth responses of Quercus castaneifolia seedlings from five different provenances from west to east of the Hyrcanian forest were investigated along their ecological gradients (from high to low precipitation). An experimental design was executed under controlled conditions at eight different irradiance levels (10, 20, 30, 40, 50, 60, 70 and 100 % full irradiance). Results showed that the irradiance is probably the most important determinant of variation in seedling characteristics. Among all investigated variables, variability in seedling size was affected significantly by provenance, while seedling morphology and their architectural response was affected by different levels of irradiance in a curvilinear manner. The biggest changes were observed at lowest irradiance levels (10–20 %) while at higher irradiance (70–100 %) the curves flatten. It was shown that, unlike at low irradiance levels, there is little capacity in seedling morphology to acclimatize with high irradiance intensity. Attaining maximal biomass varies across provenances and irradiance gradient. The highest biomass for the five provenances could be ranked as follows: 20–60 % and 50–60 % for the wetter and drier provenances, respectively. These results demonstrated that the light requirement increases from wetter to drier provenances, with a negative relationship between light requirement and precipitation regime. Different responses to irradiance levels may be the result of genetic adaptation to the ecological conditions prevailing in native habitat, especially precipitation regime.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4


  1. Ackerly D (1997) Allocation, leaf display, and growth in fluctuating irradiance environments. In: Bazzaz FA, Grace J (eds) Plant resource allocation. Academic, San Diego, pp 231–264

  2. Akhani H, Djamali M, Ghorbanalizadeh A, Ramezani E (2010) Plant biodiversity of Hyrcanian relict forests, N Iran: an overview of the flora, vegetation, palaeoecology and conservation. Pak J Bot 42:231–258

  3. Antúnez I, Retamosa EC, Villar R (2001) Relative growth rate in phylogenetically related deciduous and evergreen woody species. Oecologia 128:172–180

  4. Baltzer JA, Thomas SC (2007) Determinants of whole-plant irradiance requirements in Bornean rain forest tree saplings. J Ecol 95:1208–1221

  5. Bloor JMG, Grubb PJ (2004) Morphological plasticity of shade-tolerant tropical rainforest tree seedlings exposed to irradiance changes. Funct Ecol 18:337–348

  6. Bonito A, Varone L, Gratani L (2011) Relationship between acorn size and seedling morphological and physiological traits of Quercus ilex L. from different climates. Photosynthetica 49:75–86

  7. Cai ZQ, Rijkers T, Bongers F (2005) Photosynthetic acclimation to irradiance changes in tropical monsoon forest woody species differing in adult stature. Tree Physiol 25:1023–1031

  8. Cardillo E, Bernal CJ (2006) Morphological response and growth of cork oak (Quercus suber L.) seedlings at different shade levels. For Ecol Manag 222:296–301

  9. Castro-Diez P, Navarro J, Pintado A, Sancho LG, Maestro M (2006) Interactive effects of shade and irrigation on the performance of seedlings of three Mediterranean Quercus species. Tree Physiol 26:389–400

  10. Chazdon RL (1988) Sunflecks in the forest understory. Adv Ecol Res 18:1–63

  11. Davi H, Barbaroux C, Dufrêne E, Françoisa C, Montpied P, Bréda N (2008) Modelling leaf mass per area in forest canopy as affected by prevailing radiation conditions. Ecol Model 211:339–349

  12. Delagrange S (2011) Irradiance- and seasonal-induced plasticity in leaf morphology, N partitioning and photosynthetic capacity of two temperate deciduous species. Environ Exp Bot Paris 70:1–10

  13. Domoers M, Kaviani M, Schaefer D (1998) An analysis of regional and intra-annual precipitation variability over Iran using multivariate statistical methods. Theor Appl Climatol 61:151–159

  14. Evans JR, Poorter H (2001) Photosynthetic acclimation of plants to growth irradiance: the relative importance of specific leaf area and nitrogen partitioning in maximizing carbon gain. Plant Cell Environ 24:755–767

  15. Fila G, Sartorato I (2011) Using Leaf Mass per Area as predictor of irradiance interception and absorption in crop/weed monoculture or mixed stands. Agric For Meteorol 151:575–584

  16. Gandour M, Khouja ML, Toumi L, Triki S (2007) Morphological evaluation of cork oak (Quercus suber): mediterranean provenance variability in Tunisia. Ann For Sci 64:549–555

  17. Gratani L, Meneghini M, Pesoli P, Crescente MF (2003) Structural and functional plasticity of Quercus ilex seedlings of different provenances in Italy. Trees 17:515–521

  18. Gratani L, Convone F, Larcher W (2006) Leaf plasticity in response to irradiance of three evergreen species of the Mediterranean maquis. Trees 20:549–558

  19. Gueirn GR, Wen H, Lowe AJ (2012) Leaf morphology shift linked to climate change. Biol Lett 9:1–5

  20. Hamerlynck E, Knapp A (1996) Photosynthetic and stomatal responses to high temperature and irradiance in two oaks at the western limit of their range. Tree Physiol 16:557–565

  21. Huante P, Rincon E, Acosta I (1995) Nutrient availability and growth rate of 34 woody species from a tropical deciduous forest in Mexico. Funct Ecol 9:849–858

  22. James SA, Bell DT (2000) Influence of irradiance availability on leaf structure and growth of two Eucalyptus globulus ssp. globulus provenances. Tree Physiol 20:1007–1018

  23. Kelly J, Jose S, Nichols JD, Bristow M (2009) Growth and physiological response of six Australian rainforest tree species to a irradiance gradient. For Ecol Manag 257:287–293

  24. Kobe RK (2006) Sapling growth as a function of irradiance and landscape-level variation in soil water and foliar nitrogen in northern Michigan. Oecologia 147:119–133

  25. Oliveira G, Peñuelas J (2002) Comparative protective strategies of Cistus albidus and Quercus ilex facing photoinhibitory winter conditions. Environ Exp Bot 47:281–289

  26. Osunkoya OO, Ash JE, Hopkins MS, Graham AM (1994) Influence of seed size and seedling ecological attributes on shade-tolerance of rain-forest tree species in Northern Queensland. Int J Ecol Environ Sci 82:149–163

  27. Pearcy RW (1999) Responses of plants to heterogeneous irradiance environments. In: Pugnaire FI, Valladares F (eds) Handbook of functional plant ecology. Dekker, New York, pp 269–314

  28. Pérez-Ramos HM, Gómes-Aparicio L, Villar R, Garcia LV, Marañón T (2010) Seedling growth and morphology of three oak species along field resource gradients and seed mass variation: a seedling age-dependent response. J Veg Sci 21:419–437

  29. Poorter L (1999) Growth response of 15 rain-forest tree species to an irradiance gradient: the relative importance of morphological and physiological traits. Funct Ecol 13:396–410

  30. Puerta-Piñero C, Gómez JM, Zamora R (2006) Species-specific effects on topsoil development affect Quercus ilex seedling performance. Acta Oecologica 29(1):65-71

  31. Puerta-Piñero C, Gómez JM, Valladares F (2007) Irradiance and oak seedling survival and growth in a heterogeneous environment. For Ecol Manag 242:462–469

  32. Quero JL, Villar R, Marañón T, Zamora R, Vega D, Sack L (2008) Relating leaf photosynthetic rate to whole plant growth: drought and shade effects on seedlings of four Quercus species. Funct Ecol 35:725–737

  33. Rebbeck J, Scherzer A, Gottschalk K (2012) Do chestnut, northern red, and white oak germinant seedlings respond similarly to irradiance treatments? II. Gas exchange and chlorophyll responses. Can J For Res 42:1025–1037

  34. Reich PB, Walters MB, Ellsworth DS (1992) Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecol Monogr 62:365–392

  35. Reich PB, Walters MB, Tjoelker MG, Vanderklein D, Buschena C (1998) Photosynthesis and respiration rates depend on leaf and root morphology and nitrogen concentration in nine boreal tree species differing in relative growth rate. Funct Ecol 12:395–405

  36. Rijkers T, de Vries PJ, Pons TL, Bongers F (2000) Photosynthetic induction in saplings of three shade-tolerant tree species: comparing understory and gap habitats in a French Guiana rain forest. Oecologia 125:331–340

  37. Rouhi-moghaddam E, Hossini SM, Ebrahimi E, Tabari M, Rahmani A (2008) Comparision of growth, nutrition and soil properties of pure stand of Quercus Castaneifolia and mixed with Zelkova Carpinifolia in the hyrcanian forests of Iran. For Ecol Manag 255:1149–1160

  38. Rozendaal DMA, Hurtado VH, Poorter L (2006) Plasticity in leaf traits of 38 tropical tree species in response to irradiance; relationships with irradiance demand and adult stature. Funct Ecol 20:207–216

  39. Sabeti H (1994) Forests, trees and shrubs of Iran. Yazd University Press, Yazd

  40. Silvertown JW, Charlesworth D (2001) Introduction to plant population biology. Blackwell, New York

  41. Sugiura D, Tateno M (2011) Optimal Leaf-to-Root ratio and leaf nitrogen content determined by irradiance and nitrogen availabilities. PLoS ONE 6(7):e22236. doi:10.1371/journal.pone.0022236

  42. Thery M (2001) Forest irradiance and its influence on habitat selection. Plant Ecol 153:251–261

  43. Valladares F, Martinez-Ferri E, Balaguer L, Perez-Corona and Manrique E (2000) Low leaf-level response to irradiance and nutrients in Mediterranean evergreen oaks: a conservative resource-use strategy? New Phytol 148:79–91

  44. Veenendaal EM, Swaine MD, Lecha RT, Falsh MF, Abebrese IK, Owusu-Afriyie K (1996) Responses of West African forest tree seedlings to irradiance and soil fertility. Funct Ecol 10:501–511

  45. Xu F, Guo W, Xu W, Wang R (2008) Habitat effects on leaf morphological plasticity in Quercus acutissima. Acta Biol Cracov Bot 50:19–26

  46. Ye Z, Zhao ZA (2010) Modified rectangular hyperbola to describe the irradiance- response curve of photosynthesis of Bidense pilosa L. grown under low and high irradiance conditions. Front Agric China 4:50–55

  47. Zavala MA, Espelta JM, Retana J (2000) Constraints and trade-offs in Mediterranean plant communities. The case of Holm oak–Aleppo Pine forests. Bot Rev 66:119–149

Download references


We are grateful to Hamed Asadi for his logistic support, Mehrdad Zarafshar for taking care of the plants, and Soghra Azizi and Sodabeh Gharemahmodli for their help with harvesting. Bahram Naseri is acknowledged for his help in collecting seeds, Sattar Ezatti and Carolina Puerta-Piñero for improving the manuscript, and Ellen Vuosalo Tavakoli (University of Mazandaran) for editing the English text.

Author information

Correspondence to Seyed Gholamali Jalali.

About this article

Cite this article

Babaei Soustani, F., Jalali, S.G., Sohrabi, H. et al. Growth responses to irradiance regime along an ecological gradient of Quercus castaneifolia seedlings of different provenance. Ecol Res 29, 245–255 (2014).

Download citation


  • Light requirement
  • Morphological response
  • Temperate Hyrcanian forest
  • Site heterogeneity