Ecological Research

, Volume 25, Issue 1, pp 51–58 | Cite as

Gas exchange of juvenile and mature trees of Alnus jorullensis (Betulaceae) at sites with contrasting humidity in the Venezuelan Andes

  • Frida I. PiperEmail author
  • Lohengrin A. Cavieres
Original Article


The reduction of photosynthetic rates with tree age has been proposed as a major driver of the productivity declination along ontogeny. It is not clear, however, how environmental humidity affects stomatal conductance and biochemical potential of photosynthesis in trees belonging to different age-classes. We assessed daily cycles of gas exchange on leaves of juvenile and mature individuals of the tropical high-elevation tree Alnus jorullensis Kunth (Betulaceae), at two sites with contrasting precipitation in the Venezuelan Andes. Photosynthesis and stomatal conductance were higher in juvenile trees during the morning and at noon in the mesic site, and were in general similar between age-classes in the wet site. Under light-saturating conditions, the net photosynthetic rate was similar between the age-classes at the wet site and higher for juvenile trees at the mesic site, whereas stomatal conductance did not differ between age-classes and was higher at the wet site. Daily cycles of gas exchange and a type II regression model between photosynthesis and intercellular CO2 concentration indicated that the better performance of juvenile trees at the mesic site was due to lower non-stomatal limitations. These results support the proposal that non-stomatal limitations—rather than stomatal ones—are involved in the decay of photosynthesis in mature trees, and suggest that such limitations may be evident only under drier conditions.


Alnus High altitude Ontogeny Photosynthesis Venezuelan Andes 



We thank Fermín Rada, Marino Cabrera, Gerardo Pérez, Elysaúl Rangel, and David Martínez for their assistance in the field. We appreciated the worthy contribution of the editor and three anonymous reviewers who helped us to significantly improve the manuscript from its early version. This research was supported by the Red Latinoamericana de Botánica (RLB), fellowship P5-1993. Frida I. Piper has a postdoctoral fellowship, Fondecyt 3080057. BBVA Prize for Conservation of Biodiversity in Latin America 2004 is also acknowledged. This paper is part of the research activities of the Institute of Ecology and Biodiversity (IEB) funded by P05-002 F ICM.


  1. Ataroff M (2003) Selvas y bosques de montaña. In: Aguilera M, Azócar A, González-Jiménez E (eds) Biodiversidad en Venezuela. CONICIT-Fundación Polar, Caracas, pp 762–810Google Scholar
  2. Barnard HR, Ryan MG (2003) A test of the hydraulic limitation hypothesis in fast-growing Eucalyptus saligna. Plant Cell Environ 26:1235–1245CrossRefGoogle Scholar
  3. Becker P, Meinzer FC, Wullschleger SD (2000) Hydraulic limitation of tree height: a critique. Funct Ecol 14:4–11CrossRefGoogle Scholar
  4. Bond B (2000) Age-related changes in photosynthesis of woody plants. Trends Plant Sci 5:349–353CrossRefPubMedGoogle Scholar
  5. Cavender-Bares J, Bazzaz FA (2000) Changes in drought response strategies with ontogeny in Quercus rubra: implications for scaling from seedlings to mature trees. Oecologia 124:8–18CrossRefGoogle Scholar
  6. Chaves MM, Maroco JP, Pereira JS (2003) Understanding plant responses to drought: from genes to the whole plant. Funct Plant Biol 30:239–264CrossRefGoogle Scholar
  7. Cunningham S (2004) Stomatal sensitivity to vapour pressure deficit of temperate and tropical evergreen rainforest trees of Australia. Trees 18:399–407CrossRefGoogle Scholar
  8. Cunningham SC (2005) Photosynthetic responses to vapour pressure deficit in temperate and tropical evergreen rainforest trees of Australia. Oecologia 142:521–528CrossRefPubMedGoogle Scholar
  9. Day ME, Greenwood MS, White AS (2001) Age-related changes in foliar morphology and physiology in red spruce and their influence on declining photosynthetic rates and productivity with tree age. Tree Physiol 21:1195–1204PubMedGoogle Scholar
  10. Delzon S, Bosc A, Cantet L, Loustau D (2005) Variation of the photosynthetic capacity across a chronosequence of maritime pine correlates with needle phosphorus concentration. Ann For Sci 62:537–543CrossRefGoogle Scholar
  11. Donovan LA, Ehleringer JR (1991) Ecophysiological differences among juvenile and reproductive plants of several woody species. Oecologia 86:594–597CrossRefGoogle Scholar
  12. Falster DS, Warton DI, Wright IJ (2003) Standardised major axis tests and routines. In, 1.0 edn. Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia
  13. Hubbard RM, Bond BJ, Ryan MG (1999) Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees. Tree Physiol 19:165–172PubMedGoogle Scholar
  14. Kenzo T, Ichie T, Watanabe Y, Yoneda R, Ninomiya I, Koike T (2006) Changes in photosynthesis and leaf characteristics with tree height in five dipterocarp species in a tropical rain forest. Tree Physiol 26:865–873PubMedGoogle Scholar
  15. McDowell N, Licata J, Bond B (2005) Environmental sensitivity of gas exchange in different-sized trees. Oecologia 145:9–20CrossRefPubMedGoogle Scholar
  16. Mencuccini M et al (2005) Size-mediated ageing reduces vigour in trees. Ecol Lett 8:1183–1190CrossRefGoogle Scholar
  17. Monasterio M, Reyes S (1980a) Diversidad ambiental y variación de la vegetación en los páramos de los Andes Venezolanos. In: Monasterio M (ed) Estudios ecológicos en los páramos andinos. Ediciones de la Universidad de los Andes, Mérida, Venezuela, pp 47–92Google Scholar
  18. Monasterio M, Reyes S (1980b) Las formaciones vegetales de los páramos de Venezuela. In: Monasterio M (ed) Estudios ecológicos de los páramos andinos. Ediciones de la Universidad de los Andes, Mérida, Venezuela, pp 93–158Google Scholar
  19. Nabeshima E, Hiura T (2008) Size-dependency in hydraulic and photosynthetic properties of three Acer species having different maximum sizes. Ecol Res 23:281–288CrossRefGoogle Scholar
  20. Nieto VM, Rodriguez J (2003) Species descriptions. In: Vozzo JA (ed) Tropical tree seed manual. United States Department of Agriculture Forest Service, Washington, DC, pp 292–293Google Scholar
  21. Niinemets Ü (2002) Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris. Tree Physiol 22:515–535PubMedGoogle Scholar
  22. Ortuño M, García-Orellana Y, Conejero W, Ruiz-Sánchez M, Alarcón J, Torrecillas A (2006) Stem and leaf water potentials, gas exchange, sap flow, and trunk diameter fluctuations for detecting water stress in lemon trees. Trees Struct Funct 20:1–8Google Scholar
  23. Park SY, Furukawa A (1999) Photosynthetic and stomatal responses of two tropical and two temperate trees to atmospheric humidity. Photosynthetica 36:181–186CrossRefGoogle Scholar
  24. Pennisi E (2005) Tree growth: the sky is not the limit. Science 310:1896–1897CrossRefPubMedGoogle Scholar
  25. Phillips N, Bond BJ, McDowell NG, Ryan MG (2002) Canopy and hydraulic conductance in young, mature and old Douglas-fir trees. Tree Physiol 22:205–211PubMedGoogle Scholar
  26. Rijkers T, Pons TL, Bongers F (2000) The effect of tree height and light availability on photosynthetic leaf traits of four neotropical species differing in shade tolerance. Funct Ecol 14:77–86CrossRefGoogle Scholar
  27. Ryan MG, Yoder BJ (1997) Hydraulic limits to tree height and tree growth. BioScience 47:235–242CrossRefGoogle Scholar
  28. Ryan MG, Phillips N, Bond BJ (2006) The hydraulic limitation hypothesis revisited. Plant Cell Environ 29:367–381CrossRefPubMedGoogle Scholar
  29. Smith AP, Young TP (1987) Tropical Alpine plant ecology. Ann Rev Ecol Syst 18:137–158CrossRefGoogle Scholar
  30. Sokal RR, Rohlf FJ (1995) Biometry: the principles and practice of statistics in biological research, 3rd edn. W. H. Freeman and Company, New YorkGoogle Scholar
  31. Thomas SC, Winner WE (2002) Photosynthetic differences between saplings and adult trees: an integration of field results by meta-analysis. Tree Physiol 22:117–127PubMedGoogle Scholar
  32. Vareschi V (1970) Flora de los páramos de Venezuela. Ediciones del Rectorado, Mérida, VenezuelaGoogle Scholar
  33. Yoder BJ, Ryan MG, Waring RH, Schoettle AW, Kaufmann MR (1994) Evidence of reduced photosynthetic rates in old trees. For Sci 40:513–527Google Scholar

Copyright information

© The Ecological Society of Japan 2009

Authors and Affiliations

  1. 1.Departamento de Botánica, Facultad de Ciencias Naturales y OceanográficasUniversidad de ConcepciónConcepciónChile
  2. 2.Centro de Investigación en Ecosistemas de la Patagonia (CIEP)CoyhaiqueChile
  3. 3.Instituto de Ecología y Biodiversidad (IEB)SantiagoChile

Personalised recommendations