, Volume 23, Issue 3, pp 509–519

Physiological and morphological response to water deficit in seedlings of five provenances of Pinus canariensis: potential to detect variation in drought-tolerance

  • Rosana López
  • Jesús Rodríguez-Calcerrada
  • Luis Gil
Original Paper


To assess the potential of short-term screenings for drought resistance at the seedling stage to detect ecotypic variation and predict field performance, we studied the responses to water deficit of seedlings of Pinus canariensis from five geographic origins under controlled conditions and compared these responses with the performance of provenances in a multi-site field trial. Leaf water potential, the osmotic component, leaf chlorophyll fluorescence and growth and biomass partitioning were measured as seedlings were subjected for 11 days to two levels of osmotic potential generated by polyethylene glycol (PEG 6000), −1 MPa (slowly imposed water deficit; S) and −1.5 MPa (fast imposed water deficit; F), and a control treatment (no PEG added to the nutrient solution; C). Leaf water potential declined to final mean values of −1.2, −2.7 and −4.7 MPa in the C, S and F treatments, respectively. The ratio of variable to maximum chlorophyll fluorescence declined to final mean values of 0.77, 0.66 and 0.40 in the C, S and F treatments, respectively, with no differences amongst provenances. All provenances showed an active osmotic adjustment (OA) in response to water deficit which varied depending on the drying rate. A slow imposition of water deficit favoured solute accumulation. Pooling all treatments, the index of OA ranged from 0.28 to 0.40, but rose considerably when only C and S treatments were considered (0.56 to 0.70). There was a positive and significant correlation between the overall index of OA (all treatments pooled) and the drought period in the site of origin, suggesting ecotypic variation in OA as a result of drought duration. Seedlings allocated more dry matter to roots than shoots when subjected to moderate and slowly imposed water deficit; only one provenance showed no increase in the root to shoot ratio at the end of the treatment period compared with control seedlings. Responses to controlled water deficits were only qualitatively related to performance (survival and growth) of provenances in several field sites, indicating the involvement of complex mechanisms to cope with drought under natural conditions. However, the provenance with the highest overall index of OA outgrew and outsurvived the other provenances in the most arid site, and the only provenance not modifying the root to shoot ratio in response to water deficit survived the least in all field sites. Acclimation of root to shoot ratio and net solute accumulation to water deficit could hence favour drought-tolerance beyond the seedling stage and be used as preliminary predictors of field performance.


Drought Drying rates Osmotic adjustment Chlorophyll fluorescence Dry matter allocation Early screening 


  1. Atzmon N, Moshe Y, Schiller G (2004) Ecophysiological response to severe drought in Pinus halepensis Mill. Trees of two provenances. Plant Ecol 171:15–22. doi:10.1023/B:VEGE.0000029371.44518.38 CrossRefGoogle Scholar
  2. Baquedano FJ, Castillo FJ (2006) Comparative ecophysiological effects of drought on seedlings of the Mediterranean water-saver Pinus halepensis and water-spenders Quercus coccifera and Quercus ilex. Trees Struct Funct 20:689–700Google Scholar
  3. Baquedano FJ, Castillo FJ (2007) Drought tolerance in the Mediterranean species Quercus coccifera, Quercus ilex, Pinus halepensis, and Juniperus phoenicea. Photosynthetica 45:229–238. doi:10.1007/s11099-007-0037-x CrossRefGoogle Scholar
  4. Bloom AJ, Chapin FS, Mooney HA (1985) Resource limitation in plants-an economic analogy. Annu Rev Ecol Evol Syst 16:363–392Google Scholar
  5. Bradshaw AD (1965) Evolutionary significance of phenotypic plasticity in plants. Adv Genet 13:115–155. doi:10.1016/S0065-2660(08)60048-6 CrossRefGoogle Scholar
  6. Calamassi R, Della Rocca G, Falusi M, Paoletti E, Strati S (2001) Resistance to water stress in seedlings of eight European provenances of Pinus halepensis Mill. Ann For Sci 58:663–672. doi:10.1051/forest:2001153 CrossRefGoogle Scholar
  7. Chazen O, Hartung W, Neumann PM (1995) The different effects of PEG 6000 and NaCl on leaf development are associated with differential inhibition of root water transport. Plant Cell Environ 18:727–735. doi:10.1111/j.1365-3040.1995.tb00575.x CrossRefGoogle Scholar
  8. Clifford SC, Arndt SK, Corlett JE, Joshi S, Sankhla N, Popp M, Jones HG (1998) The role of solute accumulation, osmotic adjustment and changes in cell wall elasticity in drought tolerance in Ziziphus mauritiana (Lamk). J Exp Bot 49:967–977. doi:10.1093/jexbot/49.323.967 CrossRefGoogle Scholar
  9. Climent J, Chambel MR, López R, Mutke S, Alía R, Gil L (2006) Population divergence for heteroblasty in the Canary Island pine (Pinus canariensis, Pinaceae). Am J Bot 93:840–848. doi:10.3732/ajb.93.6.840 CrossRefGoogle Scholar
  10. Cochard H, Coll L, Le Roux X, Ameglio T (2002) Unraveling the effects of plant hydraulics on stomatal closure during water stress in walnut. Plant Physiol 128:282–290. doi:10.1104/pp.128.1.282 PubMedCrossRefGoogle Scholar
  11. Comstock J, Ehleringer J (1984) Photosynthetic responses to slowly decreasing leaf water potentials in Encelia frutescens. Oecologia 61:241–248. doi:10.1007/BF00396767 CrossRefGoogle Scholar
  12. Cregg BM, Zhang JW (2001) Physiology and morphology of Pinus sylvestris seedlings from diverse sources under cyclic drought stress. For Ecol Manage 154:131–139CrossRefGoogle Scholar
  13. Di Castri F, Goodall DW, Specht RL (1981) Ecosystems of the world 11. Mediterranean-type shrub lands. Elsevier, AmsterdamGoogle Scholar
  14. Dichio B, Xiloyannis C, Sofo A, Montanaro G (2006) Osmotic regulation in leaves and roots of olive trees during a water deficit and rewatering. Tree Physiol 26:179–185PubMedCrossRefGoogle Scholar
  15. Epron D, Dreyer E (1996) Starch and soluble carbohydrates in leaves of water-stressed oak saplings. Ann For Sci 53:263–268. doi:10.1051/forest:19960209 CrossRefGoogle Scholar
  16. Epron D, Dreyer E, Breda N (1992) Photosynthesis of oak trees (Quercus petraea (Matt) Liebl) during drought under field conditions—diurnal course of net CO2 assimilation and photochemical efficiency of photosystem-II. Plant Cell Environ 15:809–820. doi:10.1111/j.1365-3040.1992.tb02148.x CrossRefGoogle Scholar
  17. Fan S, Blake TJ (1997) Comparison of polyethylene glycol 3350 induced osmotic stress and soil drying for drought simulation in three woody species. Trees Struct Funct 11:342–348Google Scholar
  18. Fan S, Blake TJ, Blumwald E (1994) The relative contribution of elastic and osmotic adjustments to turgor maintenance of woody species. Physiol Plant 90:408–413. doi:10.1111/j.1399-3054.1994.tb00406.x CrossRefGoogle Scholar
  19. Fernández M, Gil L, Pardos JA (1999) Response of Pinus pinaster Ait. Provenances at early age to water supply. I. Water relation parameters. Ann For Sci 56:179–187. doi:10.1051/forest:19990209 CrossRefGoogle Scholar
  20. Flexas J, Ribas-Carbo M, Bota J, Galmes J, Henkle M, Martinez-Canellas S, Medrano H (2006) Decreased Rubisco activity during water stress is not induced by decreased relative water content but related to conditions of low stomatal conductance and chloroplast CO2 concentration. New Phytol 172:73–82. doi:10.1111/j.1469-8137.2006.01794.x PubMedCrossRefGoogle Scholar
  21. Gibeaut DM, Hulett J, Cramer GR, Seemann JR (1997) Maximal biomass of Arabidopsis thaliana using a simple, low-maintenance hydroponic method and favorable environmental conditions. Plant Physiol 115:317–319. doi:10.1104/pp.115.2.317 PubMedCrossRefGoogle Scholar
  22. Gieger T, Leuschner C (2004) Altitudinal change in needle water relations of Pinus canariensis and possible evidence of a drought-induced alpine timberline on Mt. Teide, Tenerife. Flora 199:100–109Google Scholar
  23. Hsiao TC, Acevedo E, Fereres E, Henderson DW (1976) Stress metabolism: water stress, growth, and osmotic adjustment. Philos Trans R Soc Lond Sev B 273:479–500. doi:10.1098/rstb.1976.0026 CrossRefGoogle Scholar
  24. Jiménez MS, Luis VC, Peters J, González-Rodríguez AM, Morales D (2005) Ecophysiological studies on Pinus canariensis. Phyton Ann Rei Bot A 45:169–177Google Scholar
  25. Jones MM, Rawson HM (1979) Influence of rate of development of leaf water deficit upon photosynthesis, leaf conductance, water use efficiency, and osmotic potential in sorghum. Physiol Plant 45:103–111. doi:10.1111/j.1399-3054.1979.tb01672.x CrossRefGoogle Scholar
  26. Jones MM, Turner NC (1980) Osmotic adjustment in expanding and fully expanded leaves of sunflower in response to water deficits. Aust J Plant Physiol 7:181–192CrossRefGoogle Scholar
  27. Kozlowski TT, Pallardy SG (2002) Acclimation and adaptive responses of woody plants to environmental stresses. Bot Rev 68:270–334. doi:10.1663/0006-8101(2002)068[0270:AAAROW]2.0.CO;2 CrossRefGoogle Scholar
  28. Lee CS, Kim JH, Yi H, You YH (2004) Seedling establishment and regeneration of Korean red pine (Pinus densiflora S. et Z.) forest in Korea in relation to soil moisture. For Ecol Manage 199:423–432Google Scholar
  29. Lilley JM, Ludlow MM (1996) Expression of osmotic adjustment and dehydration tolerance in diverse rice lines. Field Crops Res 48:185–197. doi:10.1016/S0378-4290(96)00045-7 CrossRefGoogle Scholar
  30. López R, Zehavi A, Climent J, Gil L (2007) Contrasting ecotypic differentiation for growth and survival in Pinus canariensis. Aust J Bot 55:759–769. doi:10.1071/BT07016 CrossRefGoogle Scholar
  31. Ma QF, Turner DW, Levy D, Cowling WA (2004) Solute accumulation and osmotic adjustment in leaves of Brassica oilseeds in response to soil water deficit. Aust J Agric Res 55:939–945. doi:10.1071/AR03183 CrossRefGoogle Scholar
  32. Martínez-Ferri E, Balaguer L, Valladares F, Chico JM, Manrique E (2000) Energy dissipation in drought-avoiding and drought-tolerant tree species at midday during the Mediterranean summer. Tree Physiol 20:131–138PubMedGoogle Scholar
  33. Maxwell K, Johnson GN (2000) Chlorophyll fluorescence—a practical guide. J Exp Bot 51:659–668. doi:10.1093/jexbot/51.345.659 PubMedCrossRefGoogle Scholar
  34. Mediavilla S, Escudero A (2004) Stomatal responses to drought of mature trees and seedlings of two co-occurring Mediterranean oaks. For Ecol Manage 187:281–294CrossRefGoogle Scholar
  35. Morgan JM (1992) Osmotic components and properties associated with genotype differences in osmoregulation in wheat. Aust J Plant Physiol 19:67–76Google Scholar
  36. Morgan JM, Rodríguez-Maribona B, Knights EJ (1991) Adaptation to water-deficit in chickpea breeding lines by osmoregulation: relationship to grain-yields in the field. Field Crops Res 27:61–70. doi:10.1016/0378-4290(91)90022-N CrossRefGoogle Scholar
  37. Munns R (1988) Why measure osmotic adjustment? Aust J Plant Physiol 15:717–726Google Scholar
  38. Nguyen A, Lamant A (1989) Variation in growth and osmotic regulation of roots of water-stressed maritime pine (Pinus pinaster Ait.) provenances. Tree Physiol 5:123–133PubMedGoogle Scholar
  39. Nguyen-Queyrens A, Bouchet-Lannat F (2003) Osmotic adjustment in three-year-old seedlings of five provenances of maritime pine (Pinus pinaster) in response to drought. Tree Physiol 23:397–404PubMedGoogle Scholar
  40. Parker J (1952) Desiccation in conifer leaves: anatomical changes and determination of the lethal level. Bot Gaz 14:189–198. doi:10.1086/335761 CrossRefGoogle Scholar
  41. Rodríguez-Maribona B, Tenorio JL, Conde J, Ayerve L (1992) Correlation between yield and osmotic adjustment of peas (Pisum sativum L.) under drought stress. Field Crops Res 29:15–22. doi:10.1016/0378-4290(92)90072-H CrossRefGoogle Scholar
  42. Saito T, Terashima I (2004) Reversible decreases in the bulk elastic modulus of mature leaves of deciduous Quercus species subjected to two drought treatments. Plant Cell Environ 27:863–875. doi:10.1111/j.1365-3040.2004.01192.x CrossRefGoogle Scholar
  43. Serrano L, Penuelas J, Ogaya R, Save R (2005) Tissue-water relations of two co-occurring evergreen Mediterranean species in response to seasonal and experimental drought conditions. J Plant Res 118:263–269. doi:10.1007/s10265-005-0220-8 PubMedCrossRefGoogle Scholar
  44. Tschaplinski TJ, Gebre GM, Shirshac TL (1998) Osmotic potential of several hardwood species as affected by manipulation of through fall precipitation in an upland oak forest during a dry year. Tree Physiol 18:291–298PubMedGoogle Scholar
  45. Turner DW (2006) An index of osmotic adjustment that allows comparison of its magnitude across species and experiments. Physiol Plant 127:478–482. doi:10.1111/j.1399-3054.2006.00735.x CrossRefGoogle Scholar
  46. Villar-Salvador P, Ocaña L, Peñuelas J, Carrasco I (1999) Effect of water stress conditioning on the water relations, root growth capacity, and the nitrogen and non-structural carbohydrate concentration of Pinus halepensis Mill. (Aleppo pine) seedlings. Ann Sci 56:459–465. doi:10.1051/forest:19990602 CrossRefGoogle Scholar
  47. Walter H, Lieth H (1960) Klimadiagramm-Weltatlas. Gustav Fisher Verlag, ViennaGoogle Scholar
  48. Warwick NWM, Thukten (2006) Water relations of phyllodinous and non-phyllodinous Acacias, with particular reference to osmotic adjustment. Physiol Plant 127:393–403. doi:10.1111/j.1399-3054.2006.00663.x

Copyright information

© Springer-Verlag 2008

Authors and Affiliations

  • Rosana López
    • 1
  • Jesús Rodríguez-Calcerrada
    • 1
  • Luis Gil
    • 1
  1. 1.U. D. Anatomía, Fisiología y Genética vegetal. ETSI MontesUniversidad Politécnica de Madrid. Ciudad UniversitariaMadridSpain

Personalised recommendations