, Volume 96, Issue 2, pp 219–231 | Cite as

Relative growth rate in relation to physiological and morphological traits for northern hardwood tree seedlings: species, light environment and ontogenetic considerations

  • M. B. Walters
  • E. L. Kruger
  • P. B. Reich
Original Papers


The influence of ontogeny, light environment and species on relationships of relative growth rate (RGR) to physiological and morphological traits were examined for first-year northern hardwood tree seedlings. Three Betulaceae species (Betula papyrifera, Betula alleghaniensis and Ostrya virginiana) were grown in high and low light and Quercus rubra and Acer saccharum were grown only in high light. Plant traits were determined at four ages: 41, 62, 83 and 104 days after germination. In high light (610 μmol m−2 s−1 PPFD), across species and ages, RGR was positively related to the proportion of the plant in leaves (leaf weight ratio, LWR; leaf area ratio, LAR), in situ rates of average canopy net photosynthesis (A) per unit mass (Amass) and per unit area (Aarea), and rates of leaf, stem and root respiration. In low light (127 μmol m−2 s−1 PPFD), RGR was not correlated with Amass and Aarea whereas RGR was positively correlated with LAR, LWR, and rates of root and stem respiration. RGR was negatively correlated with leaf mass per area in both high and low light. Across light levels, relationships of CO2 exchange and morphological characteristics with RGR were generally weaker than within light environments. Moreover, relationships were weaker for plant parameters containing a leaf area component (leaf mass per area, LAR and Aarea), than those that were solely mass-based (respiration rates, LWR and Amass). Across light environments, parameters incorporating the proportion of the plant in leaves and rates of photosynthesis explained a greater amount of variation in RGR (e.g. LWR*Amass, R2=0.64) than did any single parameter related to whole-plant carbon gain. RGR generally declined with age and mass, which were used as scalars of ontogeny. LWR (and LAR) also declined for seven of the eight species-light treatments and A declined in four of the five species in high light. Decreasing LWR and A with ontogeny may have been partially responsible for decreasing RGR. Declines in RGR were not due to increased respiration resulting from an increase in the proportion of solely respiring tissue (roots and stems). In general, although LWR declined with ontogeny, specific rates of leaf, stem, and root respiration also decreased. The net result was that whole-plant respiration rates per unit leaf mass decreased for all eight treatments. Identifying the major determinants of variation in growth (e.g. LWR*Amass) across light environments, species and ontogeny contributes to the establishment of a framework for exploring limits to productivity and the nature of ecological success as measured by growth. The generality of these relationships both across the sources of variation we explored here and across other sources of variation in RGR needs further study.

Key words

Relative growth rate Leaf allocation Photosynthesis Respiration Ontogeny 


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  1. Azcón-Bieto J, Osmond CB (1983) Relationship between photosynthesis and respiration. The effect of carbohydrate status on the rate of CO2 production by respiration in darkened and illuminated wheat leaves. Plant Physiol 71:574–581Google Scholar
  2. Bazzaz FA, Carlson RW (1982) Photosynthetic acclimation to variability in the light environment of early and late successional plants. Oecologia 54:313–316Google Scholar
  3. Björkman O (1981) Responses to different quantum flux densities. In: Lange OL, Nobel PS, Osmond CB, Ziegler H (eds) Encyclopedia of Plant Physiology: New Series, Vol 12A, Physiological Plant Ecology I: Responses to the Physical Environment. Springer Berlin Heidelberg New York pp 57–107Google Scholar
  4. Blackman VH (1919) The compound interest law and plant growth. Ann Bot 33:353–360Google Scholar
  5. Carter G, Smith WK (1988) Micro-habitat comparisons of transpiration and photosynthesis in three sub-alpine conifers. Can J Bot 66:963–969Google Scholar
  6. Chapin FS III, Bloom AJ, Field CB, Waring RH (1987) Plant responses to multiple environmental factors. Bio Sci 37:49–57Google Scholar
  7. Chazdon RL (1992) Photosynthetic plasticity of two rain forest shrubs across natural gap transects. Oecologia 92:586–595Google Scholar
  8. Chow P, Rolfe GL (1989) Carbon and hydrogen contents of shortrotation biomass of five hardwood species. Wood Fiber Sci 21:30–36Google Scholar
  9. Curtis JT (1959) The Vegetation of Wisconsin: An Ordination of Plant Communities. The University of Wisconsin Press 655 pGoogle Scholar
  10. Elberse WTH, Berendse F (1993) A comparative study of the growth and morphology of eight grass species from habitats with different nutrient availability. Funct Ecol 7:223–229Google Scholar
  11. Ellsworth DS, Reich PB (1992) Leaf mass per area, nitrogen content and photosynthetic carbon gain in Acer Saccharum seedlings in contrasting forest light environments. Funct Ecol 6:423–435Google Scholar
  12. Evans GC (1972) The Quantitative Analysis of Plant Growth. University of California Press p 734Google Scholar
  13. Evans JR (1983) Nitrogen and photosynthesis in the flag leaf of wheat (Triticum aestivum L.). Plant Physiol 72:297–302Google Scholar
  14. Fichtner K, Schulze ED (1992) The effect of nitrogen nutrition on growth and biomass partitioning of annual plants originating from habitats of different nitrogen availability. oecologia 92:236–241Google Scholar
  15. Garnier E (1990) Resource capture, biomass allocation and growth in herbaceous plants. TREE 6:126–131Google Scholar
  16. Givnish TJ (1986) Biomechanical constraints on crown geometry in forest herbs. In, Givnish TJ, ed. On the Economy of Plant Form and Function. Cambridge University Press pp 525–578Google Scholar
  17. Givnish TJ (1988) Adaptation to sun and shade: a whole-plant perspective. Aust J Plant Physiol 15:63–92Google Scholar
  18. Gleeson SK, Tilman D (1992) Plant allocation and the multiple limitation hypothesis. Am Nat 139:1322–1342Google Scholar
  19. Hunt R (1982) Plant Growth Curves. The Functional Approach to Growth Analysis. Edward Arnold, LondonGoogle Scholar
  20. Hunt R, Lloyd PS (1987) Growth and partitioning. New Phytol 106 [Suppl]: 235–249Google Scholar
  21. Javasekera R, Schleser GH (1991) Seasonal changes in organic carbon content of leaves of deciduous trees. J Plant Physiol 138:507–510Google Scholar
  22. Jurik TW (1986) Temporal and spatial patterns of specific leaf weight in successional northern hardwood tree species. Am J Bot 73:1083–1092Google Scholar
  23. Korner CH (1991) Some often overlooked plant characteristics as determinants of plant growth: a reconsideration. Funct Ecol 5:162–173Google Scholar
  24. Kotar J, Kovach JA, Locey CT (1988) Field Guide to Forest Habitat Types of Northern Wisconsin. Department of Forestry, University of Wisconsin-Madison and the Wisconsin Department of Natural ResourcesGoogle Scholar
  25. Kruger EL (1992) Survival, growth, root: shoot relations and ecophysiology of northern red oak (Quercus rubra L) and competing tree regeneration in response to fire and related disturbance in mesic forest openings. Ph. D. thesis. University of Wisconsin, MadisonGoogle Scholar
  26. Küppers M, Koch G, Mooney HA (1988) Compensating effects to growth of changes in dry matter allocation in response to variation in photosynthetic characteristics induced by photoperiod, light and nitrogen. Aust J Plant Physiol 15:287–298Google Scholar
  27. Lambers H (1985) Respiration in intact plants and tissues. In: Douce R, Day DA (eds) Encyclopedia of Plant Physiology: New Series, Vol 18, Higher Plant Cell Respiration, pp 418–474Google Scholar
  28. Lambers H, Poorter H (1992) Inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences. Adv Ecol Res 23:187–261Google Scholar
  29. Loach K (1967) Shade tolerance in tree seedlings. I. leaf photosynthesis and respiration in plants raised under artificial shade. New Phytol 66:607–621Google Scholar
  30. Loach K (1970) Shade tolerance in tree seedlings. II. growth analysis of tree seedlings raised under artificial shade. New Phytol 69:273–286Google Scholar
  31. Pearcy RW (1987) Photosynthetic gas exchange responses of Australian tropical forest trees in canopy, gap and understory micro-environments. Funct Ecol 1:169–178Google Scholar
  32. Pearcy RW, Calkin H (1983) Carbon dioxide exchange of C3 and C4 tree species in the understory of a Hawaiian forest. Oecologia 58:26–32Google Scholar
  33. Pfitsch WA, Pearcy RW (1989) Daily carbon gain by Adenocaulon bicolor (Asteraceae), a redwood forest understory herb, in relation to its light environment. Oecologia 80:465–470Google Scholar
  34. Pompa J, Bongers F (1988) The effect of canopy gaps on growth and morphology of seedlings of rain forest species. Oecologia 75:625–632Google Scholar
  35. Poorter H (1989) Interspecific variation in relative growth rate: on the ecological causes and physiological consequences. In: Lambers H, Cambridge ML, Konings H, Pons TL (eds) Causes and Consequences of Variation in Growth Rate and Productivity of Higher Plants. SPB Academic Publishing bv, The Hague, The Netherlands pp 45–68Google Scholar
  36. Poorter H, Pothman P (1992) Growth and carbon economy of a fast-growing and slow-growing grass species as dependent on ontogeny. New Phytol 120:159–166Google Scholar
  37. Poorter H, Remkes C (1990) Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate. Oecologia 83:553–559Google Scholar
  38. Poorter H, Remkes C, Lambers H (1990) Carbon and nitrogen economy of 24 witd species differig in relative growth rate. Plant Physiol 94:621–627Google Scholar
  39. Poorter H, Werf A van der, Atkin OK, Lambers H (1991) Respiratory energy requirements of roots vary with the potential growth rate of a plant species. Physiol Plant 83:469–475Google Scholar
  40. Reich PB, Uhl C, Walters MB, Ellsworth DS (1991a) Leaf lifespan as a determinant of leaf structure and function among 23 amazonian tree species. Oecologia 86:16–24Google Scholar
  41. Reich PB, Walters MB, Ellsworth DS (1991b) Leaf age and season influence the relationship between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees. Plant, Cell Environ 14:251–259Google Scholar
  42. Reich PB, Walters MB, Ellsworth DS (1992) Leaf lifespan in relation to leaf plant and stand characteristics among diverse ecosystems. Ecol Monogr 62(3): 365–392Google Scholar
  43. Rice SA, Bazzaz FA (1989) Quantification of plasticity of plant traits in response to light intensity: comparing phenotypes at a common weight. Oecologia 78:502–507Google Scholar
  44. SAS Institute (1985) SAS User's Guide: Statistics. Version 5, Joyner SP (ed) SAS Institute, Cary, North Carolina, USAGoogle Scholar
  45. Seeman JR, Sharkey TD, Wang JL, Osmond CB (1987) Environmental effects on photosynthesis, nitrogen-use efficency and metabolite pools in leaves of sun and shade plants. Plant Physiol 84:796–802Google Scholar
  46. Shipley B, Peters RH (1990) A test of the Tilman model of plant strategies: relative growth rate and biomass partitioning. Am Nat 136:139–153Google Scholar
  47. Strauss-Debendetti S, Bazzaz FA (1991) Plasticity and acclimation to light in tropical Moraceae of different successional position. Oecologia 87:377–387Google Scholar
  48. Turnbull H (1991) The effect of light quantity and quality during development on the photosynthetic characteristics of six Australian rainforest tree species. Oecologia 87:110–117Google Scholar
  49. Veen BW (1981) Relation between root respiration and root activity. Plant Soil 63:73–76Google Scholar
  50. Venus JC, Causton PA (1981) The biometry of plant growth. Edward Arnold, LondonGoogle Scholar
  51. Vitousek PM, Matson PA, Van Cleve K (1989) Nitrogen availability and nitrification during succession: primary, secondary, and old-field seres. In: Clarholm M, Bergstrom L, (eds) Ecology of Arable Land. Kluwer Academic, Amsterdam. pp 161–171Google Scholar
  52. Walters MB, Field CB (1987) Photosynthetic light acclimation in two rainforest Piper species with different ecological amplitudes. Oecologia 72:449–456Google Scholar
  53. Walters MB, Reich PB (1989) Response of Ulmus americana seedlings to varying nitrogen and water status. I. photosynthesis and growth. Tree Physiol 2:159–172Google Scholar
  54. Walters MB, Kruger EL, Reich PB (1993) Growth, biomass distribution and CO2 exchange of northern hardwood seedlings in high and low light: relationships with successional status and shade tolerance. Oecologia 94:7–16Google Scholar
  55. Zelitch I (1982) The close relationship between net photosynthesis and crop yield. Biosci 32:796–802Google Scholar

Copyright information

© Springer-Verlag 1993

Authors and Affiliations

  • M. B. Walters
    • 1
  • E. L. Kruger
    • 2
  • P. B. Reich
    • 1
  1. 1.Department of Forest ResourcesUniversity of MinnesotaSt. PaulUSA
  2. 2.Department of ForestryUniversity of WisconsinMadisonUSA

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