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Crown hollowing as a consequence of early shedding of leaves and shoots

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Leafing pattern has long been considered as an important element characterizing the growth strategy of tree species; however, the consequences of leafing pattern for tree-crown formation have not been fully understood. To address this issue, the dynamic events (growth, birth, and death) of current-year shoots and leaves were investigated together with their location in saplings of a pioneer tree, Alnus sieboldiana. The leafing pattern was characterized by successive emergence and shedding of short-lived leaves. The combination of successive leafing and within-crown variation in leaf production brought about characteristic outcomes in crown morphology. In the outer crown, because of continuous leaf production, the shoots achieved great extension and enormous daughter shoot production, resulting in rapid expansion of the crown. In contrast, in the inner crown, due to early termination of leaf production, the shoots completely lost their leaves early in the growing season and consequently themselves died and were shed within the season. Such quick shedding of shoots caused “crown hollowing”, i.e., the interior crown consisted of primary branches with little secondary development or foliage. These dynamic features are an effective adaptive strategy in early succession but also may be a disadvantage to maintaining foliage for longer period. Crown maintenance associated with the longevity of structural components is thought to play an important role in survival strategy of tree species.

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  1. Ackerly DD (1996) Canopy structure and dynamics: integration of growth processes in tropical pioneer trees. In: Mulkey SS, Chazdon RL, Smith AP (eds) Tropical forest plant ecophysiology. Chapman & Hall, London, pp 619–658

  2. Ackerly DD, Bazzaz FA (1995) Seedling crown orientation and interception of diffuse radiation in tropical forest gaps. Ecology 76:1134–1146. doi:10.2307/1940921

  3. Ali SM, Kikuzawa K (2005) Plasticity in leaf-area density within the crown of Aucuba japonica growing under different light levels. J Plant Res 118:307–316. doi:10.1007/s10265-005-0222-6

  4. Bazzaz FA (1979) The physiological ecology of plant succession. Annu Rev Ecol Syst 10:351–371. doi:10.1146/annurev.es.10.110179.002031

  5. Boojh R, Ramakrishnan PS (1982a) Growth strategy of trees related to successional status. 1. Architecture and extension growth. For Ecol Manage 4:359–374. doi:10.1016/0378-1127(82)90035-4

  6. Boojh R, Ramakrishnan PS (1982b) Growth strategy of trees related to successional status. 2. Leaf dynamics. For Ecol Manage 4:375–386. doi:10.1016/0378-1127(82)90036-6

  7. Chabot BF, Hicks DJ (1982) The ecology of leaf life spans. Annu Rev Ecol Syst 13:229–259. doi:10.1146/annurev.es.13.110182.001305

  8. Fisher JB (1984) Tree architecture: relationships between structure and function. In: White RA, Dickson WC (eds) Contemporary problems in plant anatomy. Academic Press, New York, pp 541–589

  9. Hallé F, Oldeman RAA, Tomlinson PB (1978) Tropical trees and forests: an architectural analysis. Springer, Berlin Heidelberg New York

  10. Hikosaka K (2005) Leaf canopy as a dynamic system: ecophysiology and optimality in leaf turnover. Ann Bot (Lond) 95:521–533. doi:10.1093/aob/mci050

  11. Horn HS (1971) The adaptive geometry of trees. Princeton University Press, Princeton

  12. Ishihara M, Kikuzawa K (2004) Species-specific variation in shoot production patterns of five birch species with respect to vegetative and reproductive shoots. Can J Bot 892:1393–1401. doi:10.1139/b04-099

  13. Ishii H, Ford ED, Dinnie CE (2002) The role of epicormic shoot production in maintaining foliage in old Pseudotsuga menziesii (Douglas-fir) trees II Basal reiteration from older branch axes. Can J Bot 80:916–926. doi:10.1139/b02-080

  14. Jones M, Harper JL (1987a) The influence of neighbours on the growth of trees. 1. The demography of buds in Betula pendula. Proc R Soc Lond B Biol Sci 232:1–18

  15. Jones M, Harper JL (1987b) The influence of neighbours on the growth of trees. 2. The fate of buds on long shoots and short shoots in Betula pendula. Proc R Soc Lond B Biol Sci 232:19–33

  16. Kikuzawa K (1983) Leaf survival of woody plants in deciduous broad-leaved forests. 1. Tall trees. Can J Bot 61:2133–2139

  17. Kikuzawa K (2003) Phenological and morphological adaptations to the light environment in two woody and two herbaceous plant species. Funct Ecol 17:29–38. doi:10.1046/j.1365-2435.2003.00707.x

  18. King DA (1990) Allometry of saplings and understory trees of a Panamanian forest. Funct Ecol 4:27–32. doi:10.2307/2389648

  19. King DA (1994) Influence of light level on the growth and morphology of saplings in a Panamanian forest. Am J Bot 81:948–957. doi:10.2307/2445287

  20. Kohyama T (1987) Significance of architecture and allometry in saplings. Funct Ecol 1:399–404. doi:10.2307/2389797

  21. Koike F (1989) Foliage-crown development and interaction in Quercus gilva and Q acuta. J Ecol 77:92–111. doi:10.2307/2260919

  22. Koike T (1988) Leaf structure and photosynthetic performance as related to the forest succession of deciduous broad-leaved trees. Plant Species Biol 3:77–87. doi:10.1111/j.1442-1984.1988.tb00173.x

  23. Koike T (1990) Autumn coloring, photosynthetic performance and leaf development of deciduous broad-leaved trees in relation to forest succession. Tree Physiol 7:21–32

  24. Kozlowski TT (1971) Growth and development of trees. Academic Press, New York

  25. Küppers M (1989) Ecological significance of above-ground architectural patterns in woody plants: a question of cost–benefit relationships. Trends Ecol Evol 4:375–379. doi:10.1016/0169-5347(89)90103-1

  26. Lovell PH, Lovell PJ (1985) The importance of plant form as a determining factor in competition and habitat exploitation. In: White J (ed) Studies on plant demography. Academic Press, London, pp 209–221

  27. Lusk CH (2002) Leaf area accumulation helps juvenile evergreen trees tolerate shade in a temperate rainforest. Oecologia 132:188–196. doi:10.1007/s00442-002-0974-9

  28. Maillette L (1982) Structural dynamics of silver birch. 1. The fates of buds. J Appl Ecol 19:203–218. doi:10.2307/2403005

  29. Pearcy RW, Valladares F (1999) Resource acquisition by plants: the role of crown architecture. In: Press MC, Scholes JD, Bacher MG (eds) Physiological plant ecology. Blackwell, London, pp 45–66

  30. Reich PB, Uhl C, Walters MB, Ellsworth DS (1991) Leaf life-span as a determinant of leaf structure and function among 23 Amazonian tree species. Oecologia 86:16–24. doi:10.1007/BF00317383

  31. Sorrensen-Cothern KA, Ford ED, Sprugel DG (1993) A model of competition incorporating plasticity through modular foliage and crown development. Ecol Monogr 63:227–304. doi:10.2307/2937102

  32. Sprugel DG, Hinckley TM, Schaap W (1991) The theory and practice of branch autonomy. Annu Rev Ecol Syst 22:309–334. doi:10.1146/annurev.es.22.110191.001521

  33. Sumida A, Komiyama A (1997) Crown spread pattern for five deciduous broad-leaved woody species: ecological significance of the retention patterns of larger branches. Ann Bot (Lond) 80:759–766. doi:10.1006/anbo.1997.0519

  34. Suzuki A (2000) Patterns of vegetative growth and reproduction in relation to branch orders: the plant as a spatially structured population. Trees (Berl) 14:329–333. doi:10.1007/s004680050226

  35. Suzuki A (2001) Resource allocation to vegetative growth and reproduction at shoot level in Eurea japonica (Theaceae): a hierarchical investment? New Phytol 152:307–312. doi:10.1046/j.0028-646X.2001.00251.x

  36. Suzuki AA (2003) Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position. Tree Physiol 23:67–71

  37. Takenaka A (1997) Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan. Tree Physiol 17:205–210

  38. Takenaka A (2000) Shoot growth responses to light microenvironment and correlative inhibition in tree seedlings under a forest canopy. Tree Physiol 20:987–991

  39. Walters MB, Reich PB (1996) Are shade tolerance, survival, and growth linked? Low light and nitrogen effects on hardwood seedlings. Ecology 77:841–853. doi:10.2307/2265505

  40. 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 94:7–16. doi:10.1007/BF00317294

  41. Wilson BF (1966) Development of the shoot system of Acer rubrum L. Harvard Forest Paper No. 14

  42. Yagi T, Kikuzawa K (1999) Patterns in size-related variations in current-year shoot structure in eight deciduous tree species. J Plant Res 112:343–352. doi:10.1007/PL00013862

  43. Yagi T (2000a) Functional differentiation of modules in broad-leaved tree species. Jpn J Ecol 50:235–250

  44. Yagi T (2000b) Morphology and biomass allocation of current-year shoots of ten tall tree species in cool temperate Japan. J Plant Res 113:171–183. doi:10.1007/PL00013928

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We thank Maki Suzuki for valuable discussions on this work. We also thank Adriano Dal Bosco and Martin J. Lechowicz for useful comments on this manuscript, Atsushi Takayanagi, Michimasa Yamasaki and other members of the Laboratory of Forest Biology, Kyoto University for helpful comments and suggestions, and Iwao Kojima and other staffs of CER for field assistance. This work was supported by the Japanese Ministry of Education and Science #11213202 and #12304047.

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Correspondence to Hiroyuki Shirakawa.

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Shirakawa, H., Kikuzawa, K. Crown hollowing as a consequence of early shedding of leaves and shoots. Ecol Res 24, 839 (2009). https://doi.org/10.1007/s11284-008-0555-4

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  • Alnus sieboldiana
  • Crown maintenance
  • Leaf longevity
  • Stem longevity
  • Successive leafing