, Volume 152, Issue 1, pp 1–12

Evidence of threshold temperatures for xylogenesis in conifers at high altitudes

  • Sergio Rossi
  • Annie Deslauriers
  • Tommaso Anfodillo
  • Vinicio Carraro


Temperature is the most important factor affecting growth at high altitudes. As trees use much of the allocated carbon gained from photosynthesis to produce branches and stems, information on the timing and dynamics of secondary wood growth is crucial to assessing temperature thresholds for xylogenesis. We have carried out histological analyses to determine cambial activity and xylem cell differentiation in conifers growing at the treeline on the eastern Alps in two sites during 2002–2004 with the aim of linking the growth process with temperature and, consequently, of defining thresholds for xylogenesis. Cambial activity occurred from May to July–August and cell differentiation from May–June to September–October. The earliest start of radial enlargement was observed in stone pine in mid-May, while Norway spruce was the last species to begin tracheid differentiation. The duration of wood formation varied from 90 to 137 days, depending on year and site, with no difference between species. Longer durations were observed in trees on the south-facing site because of the earlier onset and later ending of cell production and differentiation. The threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regressions. Xylogenesis was active when the mean daily air temperature was 5.6–8.5°C and mean stem temperature was 7.2–9°C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6–8°C that are supposed to limit growth at the treeline. Different soil temperature thresholds between sites indicated that soil temperature may not be the main factor limiting xylogenesis. This study represents the first attempt to define a threshold through comparative assessment of xylem growth and tissue temperatures in stem meristems at high altitudes.


Alps Cambial activity Cell differentiation Treeline Tree ring 


  1. Abe H, Funada R, Ohtani J, Fukazawa K (1997) Changes in the arrangement of cellulose microfibrils associated with the cessation of cell expansion in tracheids. Trees 11:328–332CrossRefGoogle Scholar
  2. Beniston M, Diaz HF, Bradley RS (1997) Climatic change at high elevation sites: an overview. Clim Change 36:233–251CrossRefGoogle Scholar
  3. Cairns DM, Malanson GP (1998) Environmental variables influencing the carbon balance at the alpine treeline: a modeling approach. J Veg Sci 9:679–692CrossRefGoogle Scholar
  4. Carrer M, Urbinati C (2001) Spatial analysis of structural and tree-ring related parameters in a timberline forest in the Italian Alps. J Veg Sci 12:643–652CrossRefGoogle Scholar
  5. Carrer M, Urbinati C (2004) Age-dependent tree-ring growth responses to climate in Larix decidua and Pinus cembra. Ecology 85:730–740Google Scholar
  6. Carrer M, Anfodillo T, Urbinati C, Carraro V (1998) High-altitude forest sensitivity to global warming: results from long-term and short-term analyses in the Eastern Italian Alps. In: Beninston M, Innes JL (eds) The impacts of climate variability on forests. Springer, Berlin Heidelberg New York, pp 171–189CrossRefGoogle Scholar
  7. Deslauriers A, Morin H (2005) Intra-annual tracheid production in balsam fir stems and the effect of meteorological variables. Trees 19:402–408CrossRefGoogle Scholar
  8. Deslauriers A, Morin H, Begin Y (2003) Cellular phenology of annual ring formation of Abies balsamea in the Quebec boreal forest (Canada). Can J For Res 33:190–200CrossRefGoogle Scholar
  9. Deslauriers A, Rossi S, Anfodillo T (2006) Dendrometer and intra-annual tree growth: what kind of information can be inferred? Dendrochronologia (in press)Google Scholar
  10. Ford ED, Robards AW, Piney MD (1978) Influence of environmental factors on cell production and differentiation in the earlywood of Picea sitchensis. Ann Bot 42:683–692Google Scholar
  11. Forster T, Schweingruber FH, Denneler B (2000) Increment puncher: a tool for extracting small cores of wood and bark from living trees. IAWA J 21:169–180Google Scholar
  12. Gindl W, Grabner M, Wimmer R (2000) The influence of temperature on latewood lignin content in treeline Norway spruce compared with maximum density and ring width. Trees 14:409–414CrossRefGoogle Scholar
  13. Grace J (1988) Temperature as a determinant of plant productivity. In: Long SP, Woodward FI (eds) Plants and temperature. Cambridge University Press, Cambridge, pp 91–107Google Scholar
  14. Grace J (1989) Tree lines. Philos Trans R Soc Lond B324:233–245Google Scholar
  15. Grace J, Norton DA (1990) Climate and growth of Pinus sylvestris at its upper altitudinal limit in Scotland: evidence from tree growth-rings. J Ecol 78:601–610CrossRefGoogle Scholar
  16. Graumlich LJ, Brubaker LB (1986) Reconstruction of annual temperature (1590–1979) for Longmire, Washington, derived from tree-rings. Quat Res 25:223–234CrossRefGoogle Scholar
  17. Gričar J, Čufar K, Oven P, Schmitt U (2005) Differentiation of terminal latewood tracheids in silver fir trees during autumn. Ann Bot 95:959–965PubMedCrossRefGoogle Scholar
  18. Hansen-Bristow K (1986) Influence of increasing elevation on growth characteristics at timberline. Can J Bot 64:2517–2523Google Scholar
  19. Hansen J, Beck E (1990) The fate and path of assimilation products in the stem of 8-year-old Scots pine (Pinus sylvestris L.) trees. Trees 4:16–21CrossRefGoogle Scholar
  20. Hansen J, Beck E (1994) Seasonal changes in the utilization and turnover of assimilation products in 8-year-old Scots pine (Pinus sylvestris L.) trees. Trees 8:172–182CrossRefGoogle Scholar
  21. Hansen J, Türk R, Vogg G, Heim R, Beck E (1997) Conifer carbohydrate physiology: updating classical views. In: Rennenberg H, Eschrich W, Ziegler H (eds) Trees: contributions to modern tree physiology. Backhuys Publishers, Leiden, pp 97–108Google Scholar
  22. Hättenschwiler S, Körner C (1995) Responses to recent climatewarming of Pinus sylvestris and Pinus cembra within their montane transition zone in the Swiss Alps. J Veg Sci 6:357–368CrossRefGoogle Scholar
  23. Holtmeier F-K (1997) Timberlines: research in Europe and North America. In: Loven L, Salmela S (eds) Pallastunturi symposium. Finnish Forest Research Institute, Finland, pp 23–36Google Scholar
  24. James JC, Grace J, Hoad SP (1994) Growth and photosynthesis of Pinus sylvestris at its altitudinal limit in Scotland. J Ecol 82:297–306CrossRefGoogle Scholar
  25. Körner C (1998) A re-assessment of high elevation treeline positions and their explanation. Oecologia 115:445–459CrossRefGoogle Scholar
  26. Körner C (2003) Alpine plant life: functional plant ecology of high mountain ecosystems, 2nd edn. Springer, Berlin Heidelberg New YorkGoogle Scholar
  27. Körner C, Paulsen J (2004) A world-wide study of high altitude treeline temperatures. J Biogeogr 31:713–732Google Scholar
  28. Kramer PJ, Kozlowski TT (1979) Physiology of woody plants. Academic, New YorkGoogle Scholar
  29. Malyshev L (1993) Levels of the upper forest boundary in northern Asia. Vegetatio 109:175–186CrossRefGoogle Scholar
  30. Motta R, Nola P (2001) Growth trends and dynamics in sub-alpine forest stands in the Varaita valley (Piedmont, Italy) and their relationships with human activities and global change. J Veg Sci 12:219–230CrossRefGoogle Scholar
  31. Oberhuber W (2004) Influence of climate on radial growth of Pinus cembra within the alpine timberline ecotone. Tree Physiol 24:291–301PubMedGoogle Scholar
  32. Oribe Y, Funada R, Kubo T (2003) Relationships between cambial activity, cell differentiation and the localisation of starch in storage tissues around the cambium in locally heated stems of Abies sachalinensis (Schmidt) Masters. Trees 17:185–192Google Scholar
  33. Philipson WR, Ward JM, Butterfield BG (1971) The vascular cambium: its development and activity. Chapman & Hall, LondonGoogle Scholar
  34. Pisaric MFJ, Holt C, Szeicz JM, Karst T, Smol JP (2003) Holocene treeline dynamics in the mountains of northeastern British Columbia, Canada, inferred from fossil pollen and stomata. Holocene 13:161–173CrossRefGoogle Scholar
  35. Potvin C, Lechowicz MJ, Tardif S (1990) The statistical analysis of ecophysiological response curves obtained from experiments involving repeated measures. Ecology 71:1389–1400CrossRefGoogle Scholar
  36. Quinn GP, Keough MJ (2002) Experimental design and data analysis for biologists. Cambridge University Press, CambridgeGoogle Scholar
  37. Rossi S, Anfodillo T, Menardi R (2006a) Trephor: a new tool for sampling microcores from tree stems. IAWA J 27:89–97Google Scholar
  38. Rossi S, Deslauriers A, Anfodillo T (2006b) Assessment of cambial activity and xylogenesis by microsampling tree species: an example at the Alpine timberline. IAWA J 27:383–394Google Scholar
  39. Rossi S, Deslauriers A, Anfodillo T, Morin H, Saracino A, Motta R, Borghetti M (2006c) Conifers in cold environments synchronize maximum growth rate of tree-ring formation with day length. New Phytol 169:279–290CrossRefGoogle Scholar
  40. SAS (1999) SAS version 8.02. SAS Institute, Cary, N.C.Google Scholar
  41. Schmitt U, Jalkanen R, Eckstein D (2004) Cambium dynamics of Pinus sylvestris and Betula spp. in the northern boreal forest in Finland. Silva Fenn 38:167–178Google Scholar
  42. Shönenberger W, Frey W (1988) Untersuchungen zur Ökologie und Technik der Hochlagenaufforstung. Forschungsergebnisse aus dem Lawinenanrissgebiet Stillberg. Schweiz Z Forstwes 139:735–820Google Scholar
  43. Smith WK, Germino MJ, Hancock TE, Johnson DM (2003) Another perspective on altitudinal limits of alpine timberlines. Tree Physiol 23:1101–1112PubMedGoogle Scholar
  44. Stevens GC, Fox JF (1991) The causes of treeline. Annu Rev Ecol Evol Syst 22:177–191CrossRefGoogle Scholar
  45. Sveinbjörnsson B (2000) North American and European treelines: external forces and internal processes controlling position. Ambio 29:388–395CrossRefGoogle Scholar
  46. Theurillat JP, Guisan A (2001) Potential impact of climate change on vegetation in the European Alps: a review. Clim Change 50:77–109CrossRefGoogle Scholar
  47. Tranquillini W (1979) Physiological ecology of the alpine timberline. Springer, Berlin Heidelberg New YorkGoogle Scholar
  48. Turner H, Streule A (1983) Wurzelwachstum und Sprossentwicklung junger Koniferen im Klimastress der alpinen Waldgrenze, mit Berücksichtigung von Mikroklima, Photosynthese und Stoffproduktion. In: Böhm W, Kutschera L, Lichtenegger E (eds) Wurzelökologie und Ihre Nutzanwendung. Irding, Gumpenstein, pp 617–635Google Scholar
  49. Uggla C, Magel E, Moritz T, Sundberg B (2001) Function and dynamics of auxin and carbohydrates during earlywood/latewood transition in Scots pine. Plant Physiol 125:2029–2039PubMedCrossRefGoogle Scholar
  50. Urbinati C, Carrer M, Sudiro S (1998) Dendroclimatic response variability of Pinus cembra L. in upper timberline forests of Italian Eastern Alps. Dendrochronologia 15:101–117Google Scholar
  51. Vaganov EA, Hughes MK, Kirdyanov AV, Schweingruber FH, Silkin PP (1999) Influence of snowfall and melt timing on tree growth in subarctic Eurasia. Nature 400:149–151CrossRefGoogle Scholar
  52. Zweifel R, Item H, Häsler R (2000) Stem radius changes and their relation to stored water in stems of young Norway spruce trees. Trees 15:50–57CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2006

Authors and Affiliations

  • Sergio Rossi
    • 1
  • Annie Deslauriers
    • 1
  • Tommaso Anfodillo
    • 1
  • Vinicio Carraro
    • 1
  1. 1.Treeline Ecology Research Unit, Dipartimento TeSAFUniversità degli Studi di PadovaLegnaroItaly

Personalised recommendations